Species diversity and ecology of Trichoptera (Caddisflies) and Plecoptera (stoneflies) in ravine ecosystems of northern ...

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Species diversity and ecology of Trichoptera (Caddisflies) and Plecoptera (stoneflies) in ravine ecosystems of northern Florida
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vii, 130 leaves : ill. ; 29 cm.
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Caddisflies -- Florida   ( lcsh )
Insects -- Ecology -- Florida   ( lcsh )
Species diversity -- Florida   ( lcsh )
Stoneflies -- Florida   ( lcsh )
Entomology and Nematology thesis, Ph. D   ( lcsh )
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Thesis (Ph. D.)--University of Florida, 2004.
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Includes bibliographical references.
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by Andrew K. Rasmussen.
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Printout.
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Vita.

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SPECIES DIVERSITY AND ECOLOGY OF TRICHOPTERA (CADDISFLIES) AND
PLECOPTERA STONEFLIESS) IN RAVINE ECOSYSTEMS OF NORTHERN
FLORIDA















By

ANDREW K. RASMUSSEN


A DISSERTATION PRESENTED TO THE GRADUATE SCHOOL
OF THE UNIVERSITY OF FLORIDA IN PARTIAL FULFILLMENT
OF THE REQUIREMENTS FOR THE DEGREE OF
DOCTOR OF PHILOSOPHY

UNIVERSITY OF FLORIDA


2004














ACKNOWLEDGMENTS

It is with great pleasure that I acknowledge the many people who are in one way or

another connected to this study. First, I would like to recognize Dr. John Capinera and the

late Dr. William "Bill" Peters for their vision and dedicated efforts that helped to affiliate

the entomology program at Florida A&M University (FAMU) with that of the University

of Florida. Dr. Capinera, in his duties as department chair, helped me enormously by

providing institutional support. I am also indebted to Debbie Hall for the excellent

administrative assistance she has offered throughout my term. I thank Dr. Sunil K.

Pancholy for providing me the office and laboratory space used to conduct my research. I

express deep gratitude to Dr. Manuel "Manny" Pescador, my graduate committee

chairman, boss, and friend, for his confidence and generous commitment of resources to

my project. Our many insect-collecting trips were both fun and productive. Appreciation

is expressed to the following past members of my committee for their participation: Dr.

Gary Buckingham (retired), Dr. Dale Habeck (retired), and Dr. William Peters

(deceased). In addition, I thank the other present members of my graduate committee for

their committee participation and thoughtful guidance: Dr. Tom Crisman, Dr. Wills

Flowers, Dr. Michael Hubbard, and Dr. John Capinera. I am grateful to the following

friends and colleagues whose company made fieldwork the most enjoyable and

rewarding part of the project: Dick Baumann, Alex Bolques, Stephanie Davis, Dana

Denson, Laurence Donelan, Wills Flowers, Steve Harris, Michael Hubbard, Jerome

Jones, Boris Kondratieff, Peter Kovarik, Corey Lewis, Charlie O'Brien, Manny Pescador,








Henry Rasmussen (my son), Don Ray, Bart Richard, Theresa Thom, and Annie Yan. I

gratefully acknowledge the following land managers and agencies for granting access to

their lands: park managers at Gold Head Branch and Torreya state parks; Greg Seamon

(The Nature Conservancy, Apalachicola Bluffs and Ravines Preserve); and Rick

McWhite, Carl Petrick, and Dennis Teague, (Natural Resources Branch, Eglin Air Force

Base). The single biggest challenge that I faced in this study was the identification of

specimens-fortunately I had the help and encouragement of excellent Trichopterists and

Plecopterists. I am deeply indebted to Steve Harris for the many ways he contributed,

including teaching me the blacklighting method used to collect adults, identifying

microcaddisfly specimens from numerous samples, and describing many of the new

species discovered during the course of the study. Other Trichopterists who generously

provided identifications and verifications were Paul Lago and Jim Glover. I express

gratitude to Dick Baumann, Stan Szczytko, and Bill Stark for identifying specimens from

some of the difficult stonefly taxa. Lastly, I express heartfelt thanks to my close friends

and family for supporting me in countless and immeasurable ways.















TABLE OF CONTENTS
page

ACKNOWLEDGMENTS....................................................... ..................

ABSTRACT ................ ...... .. ..................................................vi

CHAPTER

1 IN TRODU CTION ......................................................... ............ ....

Biogeographic Context ....... ...................... .......... ........... .......................1
Ravine Ecosystems of Northern Florida..................................................
Project Objectives and Scope................................... .......................... 14
Description of Study Areas............................................. ................... 15

2 TRICHOPTERA AND PLECOPTERA BIODIVERSITY SURVEY ................24

Previous W ork............................... .. .... .. ............ ...................... 24
M materials and M ethods ...................................... ..............................27
R results and D iscussion........................................... ............................. 30

3 ANALYSIS OF TRICHOPTERA COMMUNITY STRUCTURE AND
ENVIRONMENTAL RELATIONSHIPS................................................66

M materials and M ethods......................................................................67
Results and D iscussion.................................. ............... ...................69

4 TRICHOPTERA AND PLECOPTERA FLIGHT SEASONALITY,
AND ADULT EMERGENCE IN A RAVINE SPRINGRUN.........................82

M materials and M ethods.......................................................................82
Results and Discussion................................................ .................... 85

5 SUMMARY AND CONCLUSIONS....................................................101

R avine B iogeography.................................... ....................... ........... 101
Trichoptera and Plecoptera Species Diversity ..........................................103
Trichoptera Community Structure and Environmental Relationships..............107
Flight Seasonality and Emergence Study...............................................110
Future Research Needs ......................................... .......................... 113









page

REFERENCES CITED.......................................................................115

APPENDIX

A TRICHOPTERA DATA MATRIX......................................................1 23

B PLECOPTERA DATA MATRIX........................................................129

BIOGRAPHICAL SKETCH..................................................................130














Abstract of Dissertation Presented to the Graduate School
of the University of Florida in Partial Fulfillment of the
Requirements for the Degree of Doctor of Philosophy

SPECIES DIVERSITY AND ECOLOGY OF TRICHOPTERA (CADDISFLIES) AND
PLECOPTERA STONEFLIESS) IN RAVINE ECOSYSTEMS OF NORTHERN
FLORIDA

By

Andrew K. Rasmussen

May 2004


Chair: Manuel L. Pescador
Major Department: Entomology and Nematology

Species diversity and ecology of insects within the orders Trichoptera (caddisflies)

and Plecoptera stonefliess) were investigated in ravine streams of northern Florida.

Ravine ecosystems in Florida are known to support diverse biota, including northern

elements and endemic species, but little information concerning stream insects was

known. This study is the first to collect stream insects systematically from ravine

drainage networks across northern Florida. From West to East, the areas surveyed

included streams on Eglin Air Force Base, streams within the Apalachicola and

Ochlockonee river basins, and a stream located in peninsular Florida. Caddisfly and

stonefly species diversity was investigated at 29 stations along both upper- and lower

stream-reaches. Adult and immature insects were collected and identified, and these data

were used to characterize caddisfly and stonefly species diversity and adult flight








seasonality. Adult emergence phenology at a ravine springrun was further investigated

using emergence traps.

The survey results, based on more than 16,500 specimen identifications, document a

diverse fauna of 138 species of Trichoptera and 23 species of Plecoptera. More than 50

species are either ravine-habitat specialists, endemic to small areas of the lower Coastal

Plain, or are disjunct from northerly geographic ranges. Sixteen species recorded are

listed by the Florida Committee on Rare and Endangered Plants and Animals as

Threatened or Rare. At least 12 caddisfly species previously unknown to science were

discovered, and 11 of these have subsequently been described and named. Steephead

ravine streams in the western panhandle contain a highly endemic fauna, and the ravine

fauna of the central panhandle streams has the strongest Appalachian affinities.

In order to quantify faunal similarities among stations, cluster analysis was

performed on the survey data for macrocaddisfly species collected at 20 stations. Based

on the resultant dendogram, 5 macrocaddisfly communities were recognized among the

streams sampled. Communities appear to be hierarchically structured according to

biogeographic region at a large scale, and by habitat factors related to stream size and

ravine type at a smaller scale. Results of detrended correspondence analysis of the same

data set supported these conclusions.














CHAPTER 1
INTRODUCTION

With a budding interest in aquatic entomology and lured by the natural beauty and

mystery of ravine springruns, I undertook an investigation into the species diversity and

ecology of aquatic insects that live in these systems. After more than 10 years of

research, I am pleased to present findings concerning 2 important aquatic insect groups:

Trichoptera and Plecoptera. The following dissertation on the topic is organized by

chapters: Chapter 1 introduces the subject matter, Chapters 2, 3, and 4 address major

research objectives, and the final chapter (Chapter 5) provides a summary of the study

and conclusions. The specific objectives of this introductory chapter are to i) provide a

biogeographic context to the study; ii) present a general characterization of ravine

ecosystems in northern Florida; iii) outline the major research objectives; and iv) describe

the study areas.

Biogeographic Context

Taxa Considered

Trichoptera and Plecoptera, commonly known as caddisflies and stoneflies,

respectively, are diverse and ecologically important orders of aquatic insects. Trichoptera

are the largest order of primarily aquatic insects-globally containing over 11,000

nominal species and subspecies (Morse, 2003). In North America, more than 1300

caddisfly species have been reported (Morse, 2003), and approximately 200 of these are

known in Florida (Rasmussen, unpublished data). Plecoptera are a smaller order,








containing about 2000 described species worldwide (Stewart, 2003), approximately 600

species in North America (Stark et al., 1998) and 40 species in Florida (Rasmussen et al.,

2003).

Almost all caddisfly and stonefly species are confined to aquatic habitats during their

immature life stages (Wiggins, 1996; Stewart & Stark, 2002) and occur terrestrially only

during the relatively short-lived adult stage. Because of this aquatic dependence,

environmental conditions and the resources present in aquatic habitats are critical factors

influencing their biodiversity and evolutionary history; both groups are widely used by

environmental regulatory agencies as bioindicators to assess water quality. Caddisflies

and stoneflies are especially diverse and abundant in cool running-waters, where they are

often dominant members of benthic macroinvertebrate communities.

Much of the caddisfly and stonefly fauna found in eastern North America is endemic

to the region and has evolved in concert with eastern deciduous forests. This is

particularly true in headwater streams, where the caddisfly and stonefly faunas consist

largely of species ecologically adapted and confined to those habitats, as compared to

river faunas, which contain a higher percentage of widespread ecological generalists

(Ross, 1963). The ecology of benthic communities in headwater streams is intimately

linked with that of the terrestrial biomes in which they occur. Riparian forests, in

particular, have a great impact on the physical, chemical, and biological characteristics of

streams-the degree to which is inversely related to stream size (Vannote et al., 1980). In

northern Florida and other areas of the Southeastern Coastal Plain, many caddisfly and

stonefly taxa adapted to headwater woodland streams reach their southern range-limits








and have discontinuous peripheral populations corresponding to the patchy distribution of

equitable habitats.

Crenobiology

Streams at their source often begin as springs. The zone that includes the spring

source and downstream springrun (springbrook) is known as the crenal zone (Ward,

1992). It represents a well-bounded area with distinct biotope-the study of which is

addressed by a branch of aquatic ecology known as crenobiology. The caddisfly and

stonefly communities investigated in this study occur largely in crenal habitats; therefore,

their study falls under the discipline of crenobiology. Groundwater feeding the ravine

streams studied in this project comes from spring boils, as well as more diffuse seepage

springs along bank areas. Together, these sources produce a springrun (rheocrene).

Crenal ecosystems are attractive to ecologists because they are discrete and relatively

stable in terms of abiotic conditions at their sources, but exhibit departures along various

environmental gradients within the springrun. Crenobiological research in large artesian

springs has resulted a number of comprehensive investigations into trophic structure and

community ecology. In peninsular Florida, 2 classic studies were carried out at Silver

Springs (Odum, 1957) and Homosassa and Weekiwachee springs (Sloan, 1956). Within

temperate eastern North America, a number of influential studies have been conducted on

smaller springruns (e.g., Minckley, 1963; Minshall, 1968; Tilly, 1968). Besides

ecologists, crenal habitats are also of interest to zoogeographers because they are refugia

for relict populations of once widespread species. On the applied side, crenobiology can

be used in the biological assessment of groundwater quality. The continued interest in

crenobiology and a trend towards focusing on invertebrate biodiversity is reflected in the








papers published recently in 3 large edited-works, namely, Williams and Danks (1991);

Ferrington (1995); and Botosaneanu (1998).

Northern Florida: "Hot Spot" of Biodiversity

As a result of many interrelated factors, areas on our planet differ greatly in terms of

biodiversity. Past and present climate, geography, and geology play major roles in

shaping biomes and component biota. Availability of water, a requirement that connects

all life, is a key resource that influences the biota characteristic of a particular place.

Generally, land areas with the greatest and most unique biological diversity lie in regions

with benign climates and plentiful freshwater resources, and are geographically isolated

areas containing unique habitats with corresponding selective pressures. The panhandle

region of northern Florida satisfies all these criteria, and, not surprisingly, is a "hot spot"

of biodiversity considered to be among the top 5 regions of the continental United States

in terms of having the highest concentration of plant and animal species with restricted

distributions (Chaplin et al., 2000). Although the lower Coastal Plain occupied by

northern Florida is recognized for its rich biodiversity, there are still many invertebrate

groups for which little is known. It is within these groups that the vast majority of

unknown biodiversity remains to be discovered.

Florida's diverse natural setting has been characterized in many publications.

Excellent general accounts of climate, soils, and overviews of the many different aquatic

and marine ecosystems, as well as terrestrial upland systems in Florida, were provided in

Ecosystems of Florida (Meyers & Ewel, 1990). A comprehensive atlas summarizing

Florida's water resources is found in Femald and Purdum (1998), and surface water

resources as they relate to fisheries were covered in Seaman (1985). Papers detailing the








biodiversity of aquatic communities in Florida and the southeastern U.S. were presented

in Hackney et al. (1992). Publications focusing on the natural history of the Florida

panhandle include a general ecological characterization given by Wolfe et al. (1988), and

a thorough accounting of the natural setting and plant communities presented in Clewell

(1981).

Much of Florida's unique species diversity is documented in a series of publications

titled Rare and Endangered Biota ofFlorida. In these publications, the Florida

Committee on Rare and Endangered Plants and Animals (FCREPA) provides species lists

and documentation for species considered to be endangered, threatened, rare, or species-

of-special-concern within the state (Deyrup & Franz, 1994). As a result of new data

collected in this study, there are a number of additional species that are excellent

candidates for inclusion into future conservation listings; in Chapter 2 these species are

identified. It is through documenting and tracking species-at-risk that conservation efforts

will be most effective (Bossart & Carlton, 2002).

Historical Biogeography

The biogeographic history of Florida has been well documented. Publications

presenting general overviews included those of Neill (1957), James (1961), and Webb

(1990). Biogeographers concentrated to a great extent on Florida in attempting to

reconstruct the vegetational history of the Southeastern Coastal Plain during the

Pleistocene and Holocene. Important works included those of Delcourt and Delcourt

(1977; 1984), Watts (1980), Watts and Stuiver (1980), Watts et al. (1992), Platt and

Schwartz (1990), and Schwartz (1994). These studies support the assertion that the faunal

and floral composition of northern Florida has fluctuated throughout the Pleistocene and








Holocene in concert with climatic fluctuations and concomitant changes in temperature,

rainfall, and sea-level stands.

Major rivers draining the Southeastern Coastal Plain apparently served as important

corridors through which organisms moved in response to the climate changes

experienced during the Pleistocene and Holocene. Delcourt and Delcourt (1975; 1984)

postulated that mixed-mesophytic forest species migrated southward onto the Coastal

Plain during the late Pleistocene fostered by the moist conditions the rivers supplied to

the adjoining uplands. They concluded that dissection of these uplands into ravines with

suitable environmental conditions has resulted in the continued relict existence for many

northern plant species. By extension, this scenario can be used to account for the

distributional history of aquatic fauna associated with mixed-mesophytic forest, of which

many caddisfly and stonefly taxa are included.

It is likely that major North/South movements in biota occurred multiple times

during the various glacial and interglacial periods, suggesting that the origins of present-

day biota are multiple and complex. Hamilton and Morse (1990) investigated origins and

affinities of the caddisfly fauna of the southeastern United States and found that most

Coastal Plain endemics are more closely related to species further North and that very

few endemics have sister lineages endemic to the Coastal Plain. Hamilton and Morse

(1990, page 597) went on to infer that "the endemic Southeastern Coastal Plain caddisfly

fauna had multiple origins, perhaps the result of numerous peripheral isolations of

ancestral taxa resulting from fluctuating climates and possibly associated sea level

oscillations."









Coastal Plain Ravines as Refugia

By affording cool-adapted or stenothermal organisms the requisite environmental

conditions for survival, ravine habitats of northern Florida are present-day and historical

refugia for organisms unable to survive the harsher conditions present in predominant

habitats of the surrounding landscape. Besides containing disjunct northern elements,

ravines are also known as areas of endemism. With enough time, populations isolated

within ravine habitat-islands can speciate as a result of founder effects and local selection

pressures. The opposite phenomenon homeostasiss) can also occur, whereby ancestral

species or species representative of relict communities are preserved. In this sense,

investigations of ravine biota can be informative in reconstructing historic faunal and

floral compositions.

Neill (1957), in discussing origins of Florida's biota, references many of the early

works that describe northern Florida biota and distinctive elements of ravine habitats.

Neill (1957, page 185), noting that both northern elements and endemics are clustered

around the Apalachicola region and western panhandle, hypothesized that many of these

species were, "typical, widely ranging, Gulf Coast forms at a time when the climate was

cooler and wetter than at present." He presented as evidence for this hypothesis examples

of species that have discontinuous distributions on the Gulf Coastal Plain.

The most well known ravine-refugium within northern Florida is the Apalachicola

Bluffs and Ravines region, often referred to as "Torreya" in reference to a relict conifer,

Torreya taxifolia Am., that is largely confined to the region. In describing the area's

uniqueness, Hubbell et al. (1956, pages 20-21) stated:

Although physiographically it is no more than the dissected edge of the upland,
biotically it may be regarded as a small but distinctive natural region characterized








by its peculiar mixture of Coastal Plain and northern plants and animals, and by the
presence of a group of remarkable relic species restricted to the region. Some of its
peculiarities were pointed out by Croom as long ago as 1833-1835 and Asa Gray
(1875) made the region famous by his paper, "A Pigrimage to Torreya".

J. Speed Rogers was among the earliest zoologists to characterize streams of the Torreya

ravines and to document the uniqueness of the aquatic fauna. Rogers (1933, page 25), in

his monograph describing cranefly distributions and habitats in northern Florida, wrote of

the ravines:

Small sandy bottom brooks flow along these ravines and often pass into short
swampy reaches where they wander through tangles of standing and fallen
vegetation and over deposits of rich organic silt. Near the bottom, springs and
seepage areas are common and wet rotten wood, fungi, mosses and liverworts are
abundant.
The fauna of these ravines is as surprising and interesting as their flora, for here
a number of animals reach their southernmost limits, frequently disjunct from the
remainder of their ranges. In the Amphibia, Crustacea, Odonata, Ephemerida and
Orthoptera a number of unexpected, northern species or species with distinct
northern affinities have been discovered and among the crane-flies more than a
dozen species are found that have been taken nowhere else south of the Piedmont
region.

Ravine Ecosystems of Northern Florida

When trying to understand the diversity of organisms found in a specialized habitat

more fully, it is prudent to consider the general physical, chemical, and biological

characteristics of that habitat, as they are likely to exert considerable controls on

community composition and ecology. The following discussion provides a general

backdrop to the natural settings of the ravine ecosystems studied.

Ravine Geomorphology and Distribution

The stream systems studied occur in upland areas of northern Florida where

erosional processes have dissected the uplands to create headwater ravines and valleys in

sharp contrast to the low-relief landscapes found throughout much of the Southeastern








Coastal Plain. Ravines are steep-walled drainage networks formed both by erosion from

surface-water runoff and groundwater spring-flow. They are found in higher elevation

landforms (>50 m above mean sea level) and are often associated with river escarpments.

Topology and geomorphologic characteristics are discussed below. An account of ravine

distribution in the Florida panhandle is provided in Wolfe et al. (1988).

Steephead ravines

In landforms having high infiltration rates and large surficial-aquifer storage

capacities, as in areas with deep deposits of coarse sands, overland surface erosion is low,

and ravine formation is controlled by groundwater processes (Schumm et al., 1995).

Under these conditions, groundwater sapping causes headward erosion of overlying

deposits to form linear valleys that have steep amphitheatre-shaped heads. Ravines of this

type are known as "steepheads" and were first described by Sellards and Gunter (1918).

Steepheads are considered to be a geologic formation unique to northern Florida.

Steephead drainage networks have relatively low stream-densities (ratio of channel length

to drainage area), as compared to other drainage networks (Schumm et al., 1995), and

because of permeable soils, surface drainage lines connecting the plateau and ravine

bottom are generally absent. Steephead drainage networks tend to have a trellis pattern as

a result of tributary steepheads forming perpendicular to the main drainage axis.

Headwall slope is typically 45 degrees or greater, and depth at the head is often greater

than 30 meters.

Schumm et al. (1995) suggested that steephead advance is episodic and is promoted

by loss of vegetation on upslopes or prolonged wet periods, and that movement is halted

through groundwater capture by adjacent watersheds. Contrary to the widely accepted









fact that steephead spring-flow emerges along a relatively impermeable confining clay

layer, Schumm et al. (1995) reported finding no paleosol or hardpan in drilling at the base

of 2 steepheads on Eglin Air Force Base. This evidence and support from experimental

studies indicate that a low permeability horizon is not a requirement for steephead

formation (Schumm et al., 1995).

Steepheads in the panhandle are distributed below the Cody scarp in an east-to-west

trending line, where it appears that late Pliocene/early Pleistocene deposits of coarse

sands formed coastal sand dunes and barrier islands during higher sea level stands. Two

areas within the Florida panhandle that possess excellent examples of steephead

complexes are in the western panhandle near the coast on the western side of Eglin Air

Force Base (southern portions of Santa Rosa and Okaloosa counties) and in the central

panhandle associated with the Apalachicola River escarpment and Sweetwater Creek

(Liberty County). Steepheads occur also in several other panhandle locations (see Means,

1975), as well at a few scattered localities on upland sand ridges of the northern Florida

peninsula.

Clayhill ravines

In rolling clay hills and along river escarpments having tightly-packed upland soils,

rainfall rates often exceed infiltration rates and scouring from surface runoff results in the

formation of what I will term "clayhill" ravines, a reference to the clay content of the

soils and the name given to the associated mesic upland community-type. Another term

used to describe this type of ravine is "gully-eroded" (see Wolfe et al., 1988; Means,

2000). The drainage networks of clayhill ravines are typically dendritic and have higher

stream densities and variablity in stream flow as compared to steephead networks. Stream









flow fluctuation in these systems, however, is often moderated by groundwater flows

along ravine walls and bottoms in the form of small springs and diffuse seepage; thus,

substantial groundwater inputs result in many streams having permanent spring-fed

baseflows. Drainage lines generally originate along the upper slopes and form v-shaped

gullies. In areas of groundwater sapping, sharp vertical cuts of the overburden may also

occur. Slopes of clayhill ravine-heads and sidewalls are generally less than 30 degrees.

Clayhill ravines occur in upland landforms of the Florida panhandle above the Cody

scarp, where soils are primarily composed of Miocene-age plastics. Some of the most

well developed clayhill ravines lie within the Tallahassee Red Hills-Tifton Uplands

region in northern Gadsden and Leon counties of Florida and adjacent counties of

Georgia.

Plant and Animal Communities

The terrestrial plant and animal communities of ravine ecosystems in northern

Florida contain a diverse mix of northern, warm temperate, and endemic elements. An

accounting of this biodiversity is far from complete, and what is known is reported across

many publications. A general treatment of some of the distinctive and major components

of ravine flora and fauna is presented in Wolfe et al. (1988). Other publications dealing

with plant and animal communities of ravines include a biological survey of the

Apalachicola ravines by Leonard and Baker (1982) and a natural community survey of

Eglin Air Force Base by Kindell et al. (1997).

Mixed-species hardwoods are the most conspicuous component of ravine flora, and

these have been studied at a number of ravine systems across northern Florida (Clewell,

1981; White & Judd, 1985; Platt & Schwartz, 1990; Gibson, 1992; Kwit et al., 1998).








The very high density of different tree species occurring in ravines is accounted for by

the habitat diversity associated with gradients in elevation and soil moisture-proceeding

from xeric conditions of the ridge tops down through mesic mid-slopes to the hydric

conditions of the ravine bottom. Ravine forests typically contain oaks and hickories on

upper slopes, grading into mesic hardwoods (e.g., American beech, southern magnolia,

American holly) on midslopes, and a swamp forest community (e.g., swamp bay, sweet

bay, black gum) on hydric lower-slopes and ravine bottom. In panhandle ravines, Florida

star anise is an abundant understory species growing in saturated soils around springhead

areas. Sphagnum, other mosses, liverworts, and ferns are abundant in ravine bottoms,

which typically contain rich accumulations of leaf litter and humic soils. Much of the

faunal diversity of ravine ecosystems occurs on the forest floor and is associated with the

deep layer of leaf litter and other organic matter that serves as a food base for countless

arthropods and other invertebrates, which in turn support a far lesser number of small

vertebrates (e.g., salamanders) that feed on them.

Stream Characteristics

Because of spring-flow inputs from surficial-aquifer sources, ravine streams in

northern Florida are typically perennial and rather uniform in terms of many of their

physical and chemical characteristics. In general, the streams are clear, cool, somewhat

acidic, and low in nutrients. As compared to steephead streams, clayhill ravine streams

tend to have more variable hydroperiods and episodes of high turbidity due to greater

amounts of surface runoff and scouring after rain events. In steephead drainage networks,

the high infiltration rates and storage capacity of the surrounding landform result in

minimal runoff. Steephead springruns, over relatively short distances, can have rather








sizeable flows. Third order steephead streams discharge water at rates as high as 100 ft3

sec-'. By comparison, the baseflows of clayhill ravine streams typically are far less, even

as 5h or 6th order streams.

As water flows downstream from the ravine head, water temperature fluctuations

increase due to ambient cooling and warming. The streams with narrow- and deep

channels, typical of steephead springruns, tend to maintain more constant temperature

conditions, as compared to the shallower and less voluminous streams in clayhill regions,

which may experience significant warming in summer. High-temperature extremes can

adversely affect organisms adapted to spring-fed woodland streams and may preclude

them from living in these systems.

Benthic organisms such as caddisflies and stoneflies find a wide range of suitable

substrates in ravine streams on which to live. The stream bottom at the ravine-head

typically contains an abundance of coarse woody debris, including leaves, sticks, and

logs. Also, ravine-head mineral substrates tend to be diverse, and there often occurs pea-

gravel, pieces of clay or marls, as well as coarse sand. The stream from the ravine head

typically arises from multiple, small spring-boils and rivulets that merge to form a main

channel that then flows through the densely vegetated ravine bottom. Numerous

depositional areas of silt and organic muck are also common in the ravine heads and

upper reaches. The stream gradient is usually not great and appears to follow the slope of

the water table in most ravines systems. Debris dams can form small waterfalls in places.

Down from the ravine head, streams may meander through mesic forest or in some

systems may flow through swamp forest within a braided channel. At 3rd order and









higher, streams typically have a well-defined channel with snags and undercut banks

providing much in the way of productive benthic habitats.

Project Objectives and Scope

Given the lack of information concerning Trichoptera and Plecoptera biodiversity in

ravine ecosystems in northern Florida and the high potential for these organisms to be

important and diverse biotic components of the stream ecology, the following research

objectives were set:

i) inventory species diversity of Trichoptera and Plecoptera in ravines ecosystems of
northern Florida.

ii) characterize caddisfly and stonefly species occurrences in terms of geographic
distributional patterns, species/habitat associations, and distinctive faunal elements.

iii) investigate caddisfly community structure and distributional dynamics in relation to
geographic and habitat parameters.

iv) characterize caddisfly and stonefly flight seasonality.

v) investigate emergence of adult caddisflies and stoneflies from a ravine springrun.

This study is essentially descriptive in scope and purpose. The stream systems

studied represent several major river drainages and regions of northern Florida, and the

survey points encompass ravine streams from their upper-most headwaters through lower

stream-reaches. It is the first comprehensive study to describe and compare stonefly and

caddisfly biodiversity at several different areas on the lower Coastal Plain possessing

ravine drainage networks. I wish to stress though that faunistic studies of yet-unexplored

ravines will likely uncover important new information concerning species diversity and

community structure.









Description of Study Areas

Caddisfly and stonefly species diversity and ecology were investigated within

northern Florida at 4 different study areas (Fig. 1-1). The areas studied span much of

northern Florida and thus represent different major drainages and regions of Florida.

Three of the study areas lie within the Florida panhandle on the Eastern Gulf Coastal

Plain: the Eglin study area within the western Florida panhandle, and the Apalachicola

and Florida A&M University (FAMU) Farm study areas within the central panhandle

region. The Gold Head

study area is located on ---

the northern Florida- -

peninsula. Present ,

among the study areas .

are relatively pristine Eastern Gulf Cotal Pain

examples of both l;i ..
J, t):, < .,5i. ^ T -
steephead and clayhill / -- '
/ -,2 -. .../\
ravines from which a Eglin Study Area/ I
/ FAMUFarm j
Study Area //
wide range of different Apalachicola Study AreaGold /
Gold Head .
N Study Area ,:
stream habitats were
WE '- -
sampled. A description s
0 100 200 300 400
of the study areas and 0 M 20 30 4Kilometrs

sampling stations is
Figure 1-1. Study areas in relation to drainages and
given below. physiographic provinces of Alabama, Georgia, and
Florida.









Eglin Study Area

The Eglin study area (Fig. 1-2), largest of the 4 study areas, is located on the western

side of Eglin Air Force Base in a region where there is an extensive steephead drainage

network incised into uplands comprised largely of xeric sandhill community containing

large tracts of longleaf pine (Pinus palustris Mill.). The streams sampled feed 2 major

drainage basins, the Yellow River basin and the Choctawhatchee Bay basin. A wide array

of stream habitats is represented at the 12 stations sampled, ranging from small, ravine-

head springruns (Stations El, E3, E5) to lower stream segments with considerable

discharge (25-100 ft3 sec') (Stations E4, E8, E9, E12). Transitional in habitat between

upper steephead reaches and lower downstream reaches were middle-reach stations (E2,

E6, E7, E10, El 1). Eglin's steephead streams, because of heavy spring-inputs and

relative little surface runoff due to the highly permeable sands of the area, maintain

springrun characteristics in terms of stable flow and temperature regimes and high clarity

throughout their lengths.

At middle and lower reaches on Eglin's steephead streams, aquatic macrophytes are

particularly abundant, especially in wider stream-segments having relatively open

canopies. Some of the most common aquatic plants encountered along the clear and deep

springruns within the Eglin study area are Eriocaulon decangulare (L.), Juncus repens

(Michx.), Eleocharis spp., Orontium aquaticum (L.), Sagittaria spp., and Vallisneria

americana Michx (Theresa Thom, personal communication). Plants that are typically

emergent such as Eleocharis and Juncus repens (Michx.) often grow densely and

completely submerged in the clear waters. The freshwater red alga, Bactrachospermum

sp., is also very abundant at lower reaches.











1I
87 86045'
- 30045' 0II
0 0
a 0










E12



E10 E11




W 30030' Ealin AFB Study Area Station Localities:


I
86030'


0,0
01 2
Yi9
I >


Blackwater Bay El Okaloosa Co., Turkey Hen Ck., East Branch steephead, 0.3 km W Okaloosa Lookout
lTower off State Rd. 85. N3038'48", W8633'23".
E2 Okaloosa Co., Turkey Hen Ck., East Branch at Base Rd. 601. N30"39'27", W86"34'05".
E3 Okaloosa Co., Juniper Ck. steephead, West side of Base Rd. 231, SE of Duke Field.
N30'37'46". W86030'05".
E4 Okaloosa Co., Juniper Ck. at Base Rd. 221. N30036'28". W86031'23".
ES Okaloosa Co., Turkey Ck. Tributary, steephead East of Base Rd. 639. Choctawhatchee Bay
N3035'29", W86035'58".
E6 Okaloosa Co., Turkey Ck. at Base Rd. 637. N30'34'47", W86036'27".
0 10 E7 Okaloosa Co., Rogue Ck. at Base Rd. 625. N30033'19", W86034'51".
kilometers E8 Okaloosa Co.. Rogue Ck. at Base Rd. 233. N3033'21", W86033'44".
E9 Okaloosa Co.. Turkey Ck. at Base Rd. 232. N30033'42", W86032'10".
E10 Santa Rosa Co., Little Boiling Ck. at Base Rd. 213. N30032'29", W86051'54".
Ell Santa Rosa Co., Indigo Ck. at Base Rd. 213. N30032'27". W8650'10".
E12 Santa Rosa Co., Boiling Ck. at Base Rd. 211. N30033'55". W8652'08".




Figure 1-2. Eglin study area and locations of collecting stations. The study area is located in the western portion of Eglin Air
Force Base.












Apalachicola Study Area

The Apalachicola study area (Fig. 1-3) is located in the central Florida panhandle on

the east-side of the upper Apalachicola River in a region known as the Apalachicola

Bluffs and Ravines, sometimes referred to simply as "Torreya." The drainage network of

the region is associated with the Apalachicola/Flint river escarpment with all streams

flowing towards the lower elevations of the river's floodplain. A well-developed drainage

network comprising both steepheads and clayhill ravines has extensively dissected the

uplands to create an area with some of the sharpest relief on the Southeastern Coastal

Plain. Fortunately, a significant portion of the study area (about 8000 hectares) is

currently either within Torreya State Park or is under the stewardship of The Nature

Conservancy. Thus, conservation efforts and the prospects of preserving the area's rich

natural history are promising.

Within the study area are many small headwater streams as well as higher magnitude

(4th-6th order) streams such as Flat, Crooked, and Sweetwater creeks. Sampling stations,

representing a wide array of stream habitats, were established at 12 locations. Steephead

ravines occur predominately within the sandhill community in the southern portion of the

study area south of the main axis of Sweetwater Creek, on which station A6 is located.

Six sampling stations were within this area, including 3 upper-reach sites (A8, A9, Al 1),

2 stations further downstream on the springruns near where they enter the river floodplain

(A7, A10), and 1 station within a short isolated-ravine, very near to where its small

stream cascades into the Apalachicola River (A12). Within the more mesic clayhill-

uplands of the northern part of the study area, headwater reaches were sampled at 2

stations (A3, A5), and larger streams were sampled at 4 stations (Al, A2, A4, A6). The







19



larger streams, as a result of land/water interactions, generally have higher pH


(circumneutral) and more variable conductivity, turbidity, and temperature, as compared


to the headwater streams in which values for these parameters largely mirror spring-flow


inputs.




85 84045'


30030'


kilometers
kilometers


SGadsden County
Liberty County

Apalachicola Study Area Station Localities:
Al Gadsden Co., Flat Ck. at County Rd. 270A. N30"37'34", W8449'31".
A2 Gadsden Co., Crooked Ck. at County Rd. 270. N30034'58", W84053'02".
A3 Liberty Co., Rock Ck. Tributary. Torreya State Park, East side of Park Rd.
near Park entrance. N30033'42". W84056'48".
A4 Liberty Co., Rock Ck.. Torreya State Park. near Rock Ck. primitive campsite.
N30"34'41", W84056'15".
AS Liberty Co., Sweetwater Ck. Tributary, within The Nature Conservancy
Travelers Tract, East side of County Rd. 270, just North of Sweetwater Ck.
N30031'48", W84057'53".
A6 Liberty Co., Sweetwater Ck. at County Rd. 270. N30031'58", W84058"03".
A7 Liberty Co.. Beaver Dam Ck. with The Nature Conservancy Apalachicola Bluffs
and Ravines Preserve. N30029'13", W8459'04".
A8 Liberty Co., Little Sweetwater Ck. steephead, vicinity of Jet. State Rd. 12 and
County Rd. 270, within The Nature Conservancy Apalachicola Bluffs and
Ravines Preserve. N30028'57", W84056'27".
A9 Liberty Co., Little Sweetwater Ck. (upper end), within The Nature Conservancy
Apalachicola Bluffs and Ravines Preserve. N30"28'47". W84'57'01".
AI0 Liberty Co., Little Sweetwater Ck. (lower end), within The Nature Conservancy
Apalachicola Bluffs and Ravines Preserve. N3028'21", W8459'08".
All Liberty Co., Kelley Branch at jeep trail crossing, within The Nature Conservancy
Bluffs and Ravines Preserve. N30"28'08", W84057'51".
A12 Liberty Co., Unnamed ravine stream just North of Alum Bluff, within The
Nature Conservancy Apalachicola Bluffs and Ravines Preserve.
N30027'55", W8459'07".


Figure 1-3. Apalachicola bluffs and ravines study area and locations of collecting
stations.









FAMU Farm Study Area

The FAMU Farm study area (Fig. 1-4) is located in the central panhandle within

Ochlockonee River basin. The ravine system studied bisects property owned by Florida

A&M University, which serves as an agricultural research and extension center for the

university. The ravine is in a relatively natural state, and because of university ownership,

offers excellent opportunities for studies of the associated hardwood hammock and

spring-fed stream. Collecting

for the survey was conducted at \ .
!! |,
3 stations on this 1St-order .

stream. In addition, emergence
J-114
F I
traps were placed over the

stream at 2 locations near ,
F3
station Fl. The results of the-

emergence study are reported in "'

Chapter 4. The ravine springrun -
CONTOUR INTERVAL 10 FEET 0 1000
is a headwater for Quincy meters
FAMU Research and Extension Center (REC)
Creek, a tributary of the Little Study Area Station Localities:
F1 Gadsden Co.. Quincy Ck., headwater stream (upper reach),
River, and part of the larger FAMU REC, County Rd. 267, 7 km North of Quincy.
NRr11". u ', W84'36'50".
F2 Gadsden Co., Quincy Ck., headwater stream (middle reach),
Ochlockonee River Basin, FAMU REC. County Rd. 267, 7 km North of Quincy.
N3039' 19", W8436'51".
h d s p s of t F3 Gadsden Co., Quincy Ck., headwater stream (lower reach),
which drains portions ofthe FAMU REC, County Rd. 267, 7 km North of Quincy.
N3039' 12". W84036'56".
central Florida panhandle and
Figure 1-4. Florida A&M University Research and
southern Georgia. The ravine is Extension Center study area and locations of
collecting stations. Map taken from USGS 7.5
relatively shallow (w 15 m) as minute Dogtown quadrangle.








compared to sandhill steepheads, which can be over 30 meters deep. Within the ravine

hammock is a diverse mix of mainly hardwood species including deciduous trees such as

American beech (Fagus grandifolia Ehrh.), oaks (Quercus spp.) and hickories (Carya

spp.), as well as evergreen species including Magnolia spp., mountain laurel (Kalmia

latifolia L.) and Florida Star Anise (Illiciumfloridanum Ellis), which is the dominant

shrub species in the ravine bottom. Land use in the immediate vicinity is mainly

agricultural (planted pine and pasture) and residential.

The springrun is formed by spring flow issuing from small springs and diffuse

seepage throughout the ravine. Water is somewhat acidic (mean pH 5.6) and low in

alkalinity (, 2 mg CaCO3 L-1) with a mean conductivity of 39 pmhos cmm'. The water

temperatures in the upper sections of the springrun are relatively constant, maintaining a

temperature between 18.4 and 21.40C year-round. The springrun averages about 1 meter

in width and in most places is less than 10 cm deep; average water velocity is 0.3 m sec".

Because of shallow somewhat turbulent flows, the springrun is well oxygenated, with

oxygen levels averaging about 80% of total saturation.

Gold Head Study Area

The Gold Head study area (Fig. 1-5) is located in northeastern Florida at Mike Roess

Gold Head Branch State Park (Clay County) within the lower St Johns River basin. The

stream for which the park is named, Gold Head Branch, originates in a steephead ravine

that dissects the southeastern edge of a sand-ridge known as the Trail Ridge. The uplands

around the ravine hammock comprise primarily xeric sandhill community. White and

Judd (1985) documented an extensive flora of some 356 plant species, representing 5

communities, in the Gold Head ravine and adjacent upland. Besides Gold Head Branch,















































Figure 1-5. Gold Head Branch study area and locations of
collecting stations. Map taken from USGS 7.5 minute Gold Head
Branch quadrangle.

other waterbodies in the area include several sinkhole and sandhill lakes. Close to the

ravine head is Sheeler Lake, a deep and very old sinkhole lake (about 24,000 yrs old) that

is a well-known and important palynological site (see Watts and Stuiver, 1980). Gold

Head Branch was sampled at 2 stations, Station GI at the head of the ravine and Station








G2 near the end of the ravine just above the point at which the stream valley widens into

a delta and the stream enters a flatwoods and basin marsh before flowing into Lake

Johnson. Unlike the other ravine streams studied, Gold Head Branch terminates at a lake

and is more isolated in this sense than the other stream sites studied, which were all part

of larger stream networks. Many small spring-boils and extensive seepage occur at the

ravine head, consequently the stream rapidly gains flow within its first 50 meters. Flow

levels for Gold Head Branch, as reported in the Park Management Plan, ranged from

34,077 gallons per hour to 56,520 gallons per hour. As is typical of ravine springruns in

northern Florida, water temperature stays within a narrow range, the water is somewhat

acidic, and has low conductivity. Recent measurements taken near station G2 during

several times of year by the Florida Department of Environmental Protection ranged as

follows: pH (4.6-6.6), water temperature (17.4-23.50C), specific conductance (16-34

limhos cm-) (Lee Banks, personal communication).














CHAPTER 2
TRICHOPTERA AND PLECOPTERA BIODIVERSITY SURVEY

As noted in Chapter 1, ravine ecosystems in northern Florida have been well

documented in terms of their vegetation and geomorphology, but knowledge of ravine

faunas, especially aquatic insect groups, is relatively incomplete. This study, an

investigation into caddisfly and stonefly biodiversity, is the first in which aquatic insects

were systematically collected from headwater and downstream reaches of streams in

several upland areas of northern Florida that possess relatively pristine and exemplary

ravine ecosystems. The objectives of the survey were to inventory the species of

Trichoptera and Plecoptera occurring at the 4 study areas and to characterize species

occurrences in terms of geographic distributional patterns, species/habitat associations,

and distinctive faunal elements. Survey data presented in this chapter were also used for

analyzing Trichoptera community structure (Chapter 3) and characterizing caddisfly and

stonefly adult seasonality (Chapter 4).

Previous Work

Apalachicola Bluffs and Ravines

Of the areas studied, the Torreya ravines historically received the most attention

from early 20th century entomologists, probably a result of the region's botanical

peculiarities. Well known entomologists who studied the Torreya area's insect

biodiversity included Lewis Berner (Ephemeroptera), C.F. Byers and M.J. Westfall

(Odonata), T.H. Hubbell (Orthoptera), J. Speed Rodgers (Diptera:Tipulidae), and F.K.








Young (aquatic beetles). As indicated, Trichoptera and Plecoptera were not of primary

interest to early entomologists visiting the Torreya ravines; therefore, little effort was

devoted to their study. Results of early work on the insect groups that were studied and

other aquatic fauna such as crayfish and amphibians indicated that, like the flora, the

aquatic fauna shows close biogeographic affinities with the Appalachian highlands and

includes narrow-range endemics (Hubbell et al., 1956; Neill, 1957). Entomologists have

continued to be drawn to the Torreya area, and through their concerted collecting efforts,

much has been learned concerning the area's insect fauna, terrestrial insects in particular.

Aside from the Apalachicola ravines, other ravine habitats in northern Florida have not

been extensively sampled by entomologists-as a result, their insect fauna remains

relatively unknown.

Regional Surveys of Trichoptera and Plecoptera

In order to characterize the species composition and biodiversity of Trichoptera and

Plecoptera in ravine assemblages adequately, it was necessary to examine survey results

in the context of the component species' geographic ranges and occurrences in other

habitats. Without this broader knowledge, the fauna of a particular habitat assemblage

cannot be described in terms of its distinctiveness, and characterization must be left to

simply presenting checklists of species occurrences. Previous and ongoing studies of

caddisfly and stonefly biodiversity within northern Florida and neighboring states have

provided much of the needed information to allow placing the survey findings into a

larger context. Comprehensive inventories of the Trichoptera fauna of Alabama (Harris et

al., 1991) and Mississippi and southeastern Louisiana (Harris et al., 1982; Holzenthal et

al., 1982; Lago et al., 1982) were particularly useful in this regard.








The first statewide survey for stoneflies in Florida was a checklist of 17 species

presented in Berner (1948). The records presented in Berner's checklist were based on

specimens he collected during his expeditions in the 1930's to collect mayflies. Among

the stonefly records presented by Berner are collections from the Apalachicola ravines

region, including Torreya State Park, Sweetwater Creek and Little Sweetwater Creek.

Subsequent studies of the stonefly fauna of Florida have focused mainly on larger creeks

and rivers-particularly within the Blackwater, Ochlockonee, and Suwannee river

basins-not ravine headwaters. Recent publications that summarized the stonefly fauna

of Florida include the papers of Stark and Gaufin (1979), Pescador et al. (2000), and

Rasmussen et al. (2003); of these, the last two incorporated stonefly collection data from

this study.

Through the work of various Trichopterists, the caddisfly fauna of Florida, as with

Plecoptera, is fairly well known. Statewide survey-accounts were provided by Blickle

(1962) for Hydroptilidae and by Gordon (1984) for non-hydroptilid caddisflies. Pescador

et al. (1995) provided larval keys and an updated species-checklist with distributional

data, including preliminary data from this study, on caddisflies found within the

Apalachicola ravines and FAMU Farm ravine. Light trapping I have conducted in recent

years at various waterbodies (besides ravine streams) in central and northern Florida and

southern Georgia was complementary to this study, and these data were used to compare

ravine-assemblage species composition with those of other aquatic habitats.

Of the study areas included in this survey, the Eglin area previously was sampled the

most extensively for caddisflies. Adult caddisflies were collected from several streams on

the eastern side of Eglin Air Force Base by J.F. Scheiring. The results reported in Harris








et al. (1982) included a number of species new to science, several of which were

subsequently described in Lago and Harris (1983; 1987) and Bueno-Soria (1981). Of the

new species found on Eglin, most are narrow-range endemics-either confined entirely to

Eglin, or occupying somewhat larger ranges that extend into other parts of the western

panhandle and adjacent coastal Alabama and Mississippi. The stream reaches sampled by

Scheiring did not include steephead or upper ravine-reaches. One-time light trapping by

B.A. Armitage and M.K. Ward along several streams (downstream reaches) on the

western side of Eglin resulted in the discovery of additional new species of

microcaddisflies, subsequently described in Harris and Armitage (1987) and Harris

(1991).

Materials and Methods

Specimen collecting. Caddisfly and stonefly adults and aquatic stages from the 29

stations were sampled using a variety of qualitative collecting methods. Because less than

50% of the caddisfly and stonefly species occurring in Florida can be identified to species

in the larval stage, collection methods targeting adults were emphasized for obtaining

species-level data. For each sampling date (Table 2-1), adults, and usually aquatic stages,

were collected. Most samples were collected in the spring when most species are present

as adults. However, at stations repeatedly sampled, summer and fall samples were also

obtained so that species emerging as adults later in the year would not be missed in the

survey. Aquatic forms were collected by aquatic dipnet from various benthic

microhabitats including rootmats, aquatic macrophytes, leaf packs, snags, and mineral

substrates. Benthic sampling consisted of taking several dipnet sweeps from

representative benthic microhabitats over a 50-m stream reach. Net samples were put in








white sorting pans, and specimens were field-picked, placed in 80% ethyl alcohol, and

returned to the laboratory for identification and curation. Adult stoneflies and caddisflies

were collected primarily by light trapping, but sweep netting and a beating sheet were

also occasionally employed to supplement light-trap collections and to provide a means

of collecting stonefly and caddisfly species not readily drawn to light. The beating sheet

was most useful when temperatures were low and insects were not able to escape by

flying away. Sweep netting was done during warmer times of year to capture both insects

seen flying and to capture individuals resting on riparian vegetation. Light trapping

provided the most productive means for capturing adults of most caddisflies species, as

well as many stonefly species. Light traps each consisted of a 15-watt UV-blacklight

(BioQuip Item No. 2805) placed over a white pan (30 cm x 25 cm) containing 80%

ethyl alcohol. The collecting lights were powered by lightweight 12-volt, sealed

rechargeable-batteries (Yuasa NP7-12). Traps were placed near the water's edge and

deployed for 1.5-3 hrs beginning at dusk. Typically, 2-5 traps were run on the same night

at different stations within a study area. After trapping, the contents of the pans were

poured into 0.5-gallon plastic containers and returned to the laboratory for processing. A

total of 116 light-trap collections were made.

Specimen identification. Samples were sorted and specimens identified with the aid

of a stereomicroscope (Olympus SZH). Processing of the light-trap material often

required extensive sorting, and for samples with high numbers of non-target insects such

as moths, dipterans, beetles, etc., it was first necessary to separate them from the sample.

For most caddisfly and many stonefly species, adult males were primarily used for

making species determinations. Because many species were represented by 1 or only-a-









few individuals, samples were not sub-sampled. However, in instances of species being

represented by several hundred or more individuals in a light-trap collection, counts were

abbreviated due to time constraints. Specimen identifications were to the lowest possible

taxonomic level, and, except for some hydroptilid collections, the number of specimens

for each species record was tallied. Immature forms and females for many genera could

not be identified to species-level and were not included in the analyses. Adult

identifications usually required examination of genitalic structures. An extensive body of

taxonomic literature was consulted, and collections housed at FAMU served as reference

material. Specimens of various taxa were sent to taxonomic specialists for identifications

or verifications. Dr. Steven Harris identified all the microcaddisfly (Hydroptilidae)

collections; Dr. Paul Lago identified a large number ofhydropsychid and

polycentropodid specimens; Dr. James Glover verified identifications of Triaenodes

(Leptoceridae) species. Stonefly specialists that were consulted included Dr. Bill Stark,

Dr. Stanley Szczytko, and Dr. Richard Baumann. Synoptic voucher collections will be

deposited in the following locations: Florida A&M University Aquatic Insect Collection,

Clemson University Arthropod Collection, and in the personal collection of the author.

Descriptions of new species and deposition of type material was, or will be, detailed in

separate papers.

Data analysis. Specimen collection data from the survey were entered into a

relational database (Paradox 9.0) so that specimen data could be easily retrieved and

queried. For each species, specimen counts from collections at each sampling station

were summed and abundance data were entered into spreadsheet matrices; species

richness and abundance values were determined for the different stations and study areas.










Results and Discussion

The survey of ravines in northern Florida for Trichoptera and Plecoptera species

diversity revealed a diverse and interesting fauna. A summary of the number of species

and specimens identified from each station is presented in Table 2-1. Overall, more than

16,500 specimens were identified to species, and the 138 species of Trichoptera and the

23 species of Plecoptera identified represent approximately 70% and 55%, respectively,

of the total known faunas in Florida. The high species richness reflects the wide array of

high-quality habitats found within ravine drainage networks that span much of biota-rich

northern Florida.

Table 2-1. Summary of survey results (See Chapter 1 for explanation of station coding).
Trichoptera Plecoptera
Specimens Specimens
Station Sampling Dates Species Identified Species Identified
Code n (n) (n) (n) (n)


21.v.98; 27.x.98; 10.iii.98; 8.iv.99; 16.vi.99;
10.iv.01
28.x.98; 10.iii.99; 8.iv.99; 16.vi.99; 2.iii.00;
S11.iv.01
21.v.98; 27.x.98; 10.iii.99; 8.iv.99; 16.vi.99;
10.iv.01
19.iii.98; 21.v.98; 27.x.98; 10.iii.99; 8.iv.99;
2.iii.00; 11.iv.01
27.x.98; 7.iv.99; 16.vi.99
27.x.98; 7.iv.99; 15.vi.99
21.v.98; 28.x.98; 7.iv.99; 15.vi.99
13.xi.97
13.xi.97; 21.v.98; 10.iii.99; 7.iv.99; 15.vi.99;
1.xi.99
19.iii.98
19.iii.98
19.iii.98
18.iv.95; 7.vi.99
18.iv.95; 20.xi.98; 1.iv.99; 7.vi.99
9.iv.98; 20.xi.98; 1.iv.99; 8.vi.99
9.iv.98; 20.xi.98; 1.iv.99; 8.vi.99
1.iv.99; 8.vi.99
19.v.94; 18.iv.95; 24.vi.96; 9.iv.98; 20.xi.98;
1.iv.99; 7.vi.99
19.v.94; 7.xii.94; 22.iii.95; 30.viii.95; 26.x.95;
24.vi.96
1.iv.99; 8.vi.99
7.xii.94; 22.iii.95; 30.viii.95; 26.x.95; 24.vi.96


56 912
41 807

42 454

53 866

19 409
37 1038
56 1219
13 92
50 1680

16 100
16 121
16 124
27 375
38 343
20 126
25 277
14 38
44 965

54 697

10 84
36 298









Table 2-1. Continued.
Trichoptera Plecoptera
Specimens Specimens
Station gE Sampling Dates Species Identified Species Identified
Code C, (n) (n) (n) (n)
A10 6 7.iv.94; 19.v.94; 7.xii.94; 22.iii.95; 30.viii.95; 55 902 2 17
26.x.95
All 5 19.v.94; 22.iii.95; 30.viii.95; 26.x.95; 24.vi.96 42 378 6 14
A12 3 7.xii.94; 22.iii.95; 26.x.95 17 282 1 4
Fl 5 19.iv.94; 17.v.94; 30.iii.95; 14.ix.95; 11.xi.99 37 690 4 47
F2 5 6.v.93; 6.x.93; 17.v.94; 30.iii.95; 14.ix.95 33 270 2 2
F3 1 14.ix.95 17 34 0 0
G1 5 1.v.98; 27.vi.98; 3.x.98; 6.iii.99; 5.vi.99 28 979 2 209
G2 5 1.v.98; 27.vi.98; 3.x.98; 6.iii.99; 5.vi.99 28 1198 2 103

The following sections give an accounting of species occurrence within the study

areas, and notes are included on species geographic distribution and habitat associations;

particular attention is paid to faunal elements that are unique to, or characteristic of, the

stream habitats sampled. Species conservation status according to the Florida Committee

on Rare and Endangered Plants and Animals (FCREPA) is identified, and among the

recently discovered species, recommendations are made concerning species that should

be considered as candidates for future listing.

Faunal Elements of Special Interest

Among the stonefly and caddisfly species recorded in the survey, I recognize

approximately 50 species of special interest (Table 2-2) that represent faunal elements

distinctive of the habitats and geographic areas where they occur. Collection records for

the majority of these species suggest that the ravines surveyed contain habitats vital for

supporting local populations. These species of special interest exemplify the unique biota

found in ravine ecosystems, and their occurrences attest to the importance of ravine

habitats in supporting a distinct and diverse fauna. Species were designated as species of

special interest if they fit into one or more of the following categories: ravine crenobiont,

narrow-range endemic, disjunct, listed by FCREPA as Rare or Threatened. The









terminology "ravine crenobiont", "narrow-range endemic", and "disjunct" should be

defined to prevent misunderstanding. A species restricted to ravine-head springruns is

referred to as a "ravine crenobiont" because of its close ecological association and

confinement to ravine crenal habitats. A "narrow-range endemic" for this study is defined

as a species having its entire geographic range within the lower (coastal) areas of the

Southeastern Coastal Plain. A species referred to as "disjunct" has populations within the

study areas that are separated by considerable distances from the species' main

geographic range, typically with few, if any, intervening populations on the Southeastern

Coastal Plain.

Table 2-2. Species of special interest: ravine crenobionts, narrow-range endemics,
disjuncts, and species listed by the Florida Committee on Rare and Endangered Plants
and Animals (FCREPA) as Rare (R), Threatened (T), or of Undetermined Status (U).
Species' Study Area Ravine Narrowly Disjunct FCREPA
Occurrence Crenobiont endemic Statusb
TRICHOPTERA
Hydropsychidae
Cheumatopsyche gordonae E X T
Cheumatopsyche petersi E X R
Diplectrona modest E, A, F X
Diplectrona sp. A G X X
Philopotamidae
Chimarrafalculata E, A X
Polycentropodidae
Cernotina truncona G R
Nyctiophylax morse E X R
Polycentropus clinei G X
Polycentropusfloridensis E X T
Hydroptilidae
Hydroptila apalachicola A X
Hydroptila bribriae E X
Hydroptila circangula E X
Hydroptila eglinensis E X
Hydroptila hamiltoni E X
Hydroptila latosa E, G X R
Hydroptila lloganae E R
Hydroptila molsonae E X R
Hydroptila okaloosa E X
Hydroptila parastrepha E X
Hydroptila sarahae E X
Neotrichia armitagei E, G X
Ochrotrichia apalachicola E, A X
Orthotrichia baldufi A, F X










Table 2-2. Continued.
Speciesa Study Area Ravine Narrowly Disjunct FCREPA
Occurrence Crenobiont endemic Status
Orthotrichia curta G R
Oxyethira chrysocara G X
Oxyethira elerobi E R
Oxyethiraflorida E, G X T
Oxyethira grisea A X
Oxyethira kelleyi E X T
Oxyethira setosa A R
Beraeidae
Beraea n. sp. E X X
Brachycentridae
Micrasema n. sp. E X
Calamoceratidae
Heteroplectron americanum E, A, F X X
Lepidostomatidae
Lepidostoma griseum A X
Lepidostoma latipenne A X
Lepidostoma serratum E, F X X
Leptoceridae
Ceraclea diluta E X
Nectopsyche paludicola E X U
Oecetis daytona E, G R
Triaenodes helo A X R
Triaenodes taenia A, F X X
Molannidae
Molanna blenda E, A, F X X
Odontoceridae
Psilotretafrontalis E, A, F X X
Sericostomatidae
Agarodes logani F X X
Agarodes ziczac E X T
PLECOPTERA
Leuctridae
Leuctra cottaquilla E X
Leuctra triloba A, F X X
Perlidae
Acroneuria lycorias E, A, F X
Eccoptura xanthenes A, F X X
Perlesta sp. A E ?
Perlesta sp. B E ?
Pteronarcyidae
Pteronarcys dorsata E, A X
a Higher classification and author names are omitted (see Tables 2-3 to 2-6).
b Conservation status taken from Deyrup and Franz (1994)


Eleven species of ravine crenobionts were recorded in the survey: 9 caddisfly species

and 2 stonefly species. Of these, 3 caddisfly species (Diplectrona sp. A, Beraea n. sp.,

Agarodes logani) appear to be narrow-range endemics. The other ravine crenobionts are









species that typically occur in small, cool streams of more northern latitudes. Other

species were inventoried that are not restricted to upper reaches of ravine streams, but are

narrowly endemic or have disjunct populations occurring within the stream systems

sampled. More than 25 narrowly-endemic species were recorded from the streams

surveyed; the majority (15 species) belong to the Trichoptera family Hydroptilidae. All

study areas contained narrow-range endemics, but the Eglin study area showed by far the

highest degree of endemism. Of the 92 species of caddisflies inventoried at the Eglin

stations, 22 species (24%) are narrow-range endemics. By all indications, Eglin's spring-

fed streams, due to their high-quality habitats and physical isolation, are a major center of

caddisfly endemism within the Southeastern Coastal Plain. As discussed in the following

survey accounts, some narrow-range endemics appear to be restricted within the

particular study area, or possibly a single stream system, while other species occur across

somewhat larger areas. Differences in relative abundance were significant, and these

differences give some indication as to which species are most important to ecosystem

functioning. Many of the endemics listed appear to be not as common or as abundant

from outside the study area, and it seems probable that speciation occurred locally

autochthonouss endemics) and dispersal to a wider geographic area is limited due to

environmental factors. Because of restricted distributions of these species, they are

particularly vulnerable to extinction. Sixteen caddisfly species reported in the survey are

considered by the Florida Committee on Rare and Endangered Plants to be Rare or

Threatened (Deyrup & Franz, 1994). There were a number of species collected that are

disjunct from their main geographic ranges. Many of these species likely are unable to

tolerate high temperature extremes, and thus are restricted in Florida to thermally-








buffered steephead and clayhill ravine streams. The majority of disjuncts occur in the

central panhandle region and likely dispersed to the region from the southern

Appalachian highlands via Apalachicola-Flint-Chattahoochee watershed connections.

Survey Account of Trichoptera

Overall, 138 Trichoptera species representing 37 genera and 17 families were

identified from among the 116 samples taken at the 29 collecting stations. A total of

15,758 specimens were identified to species. The family Hydroptilidae contained the

most species (44) inventoried, followed by Leptoceridae (32), Hydropsychidae (17),

Polycentropodidae (13), Philopotamidae (5) Sericostomatidae (4), Dipeseudopsidae (3),

Brachycentridae (3), Lepidostomatidae (3), Molannidae (3), Phryganeidae (3),

Calamoceratidae (2), Limnephilidae (2), Psychomyiidae (1), Rhyacophilidae (1),

Odontoceridae (1), Beraeidae (1). Species richness for the 4 study areas was as follows:

Eglin (92 species), Apalachicola (89 species), FAMU Farm (49 species), and Gold Head

(35 species). Caddisfly faunal composition differed significantly among study areas, and

many species were recorded from only a single study area. Collections from each study

area contained the following number of species recorded solely within that particular

study area: Eglin (31 species), Apalachicola (23 species), FAMU Farm (4 species), and

Gold Head (6 species). The Trichoptera species inventory, presented in Tables 2-3 to 2-5,

is discussed below. Family subheadings and species are arranged alphabetically under

each of the 3 widely recognized Trichoptera suborders (Annulipalpia, Spicipalpia,

Integripalpia).










Table 2-3. Survey Summary (Trichoptera:Annulipalpia).
Coll. Speci Study Area (% of Total Specimens)
Species (n) mens [Collection Station Number]
(n)


Dipseudopsidae
Phylocentropus carolinus Carpenter
Phylocentropus lucidus (Hagen)
Phylocentropus placidus (Banks)
Hydropsychidae
Cheumatopsyche burksi Ross
Cheumatopsyche campyla Ross
Cheumatopsyche edista Gordon
Cheumatopsyche gordonae Lago &
Harris
Cheumatopsyche petersi Ross et al.
Cheumatopsyche pettiti (Banks)
Cheumatopsyche pinaca Ross
Cheumatopsyche virginica Denning
Diplectrona modest Banks
Diplectrona sp. A
Hydropsyche betteni Ross
Hydropsyche decalda Ross
Hydropsyche elissoma Ross
Hydropsyche incommoda Hagen
Hydropsyche rossi Flint et al.
Macrostemum carolina (Banks)
Potamyiaflava (Hagen)
Philopotamidae
Chimarra aterrima (Hagen)

Chimarrafalculata Lago & Harris
Chimarra florida Ross

Chimarra moselyi Denning
Chimarra obscure (Walker)
Polycentropodidae
Cernotina calcea Ross
Cernotina spicata Ross
Cernotina truncona Ross
Cyrnellus fraternus (Banks)
Neureclipsis crepuscularis (Walker)
Neureclipsis melco Ross
Nyctiophylax affinis (Banks)
Nyctiophylax morse Lago & Harris
Nyctiophylax serratus Lago & Harris
Polycentropus blicklei Ross &
Yamamoto
Polycentropus cinereus Hagen

Polycentropus clinei (Milne)
Polycentropusfloridensis Lago & Harris
Psychomyiidae
Lvpe diverse (Banks)


E(14)[9]; A(80)[2,5-7,10]; F(6)[1,2]
A(39)[2,3,7,9-12]; F(61)[1-3]
E(27)[1,2]; A(73)[2,6,9,12]

E(50)[2]; F(50)[2]
A(100)[1,2,4,6,10,12]
E(8)[2,9]; A(84)[1-4,6,7,10,11]; F(8)[1,2]
E(100)[1-8,10,11]

E(100)[1,2,4-7,9-12]
E(8)[2,3,7]; A(41)[2-4, 6,7,9-12]; F(51)[1,2]
A(24)[1,2,4,6-11]; F(1)[1,2]; G(75)[1,2]
E(92)[1-4,6,7,9,10,12]; A(8)[10,11]
E(57)[1-8]; A(26)[3,7-12]; F(17)[1,2]
G(100)[1]
F(100)[2,3]
G(100)[1]
E(72)[1-7,9-12]; A(27)[2,6,7,9-11]; F(1)[1,3]
E(0.2)[1,6]; A(99.7)[1,2,4-12]; F(0.1)[1]
E(0.6)[7]; A(99.4)[1,2,4,6,7,10-12]
E(100)[1-7,9]
A(100)[4,6,10,12]


27 152 E(24)[1,3,7,9]; A(31)[7,10-12]; F(4)[1,2];
G(41)[1,2]
47 472 E(89)[1-11]; A(11)[7-11]
26 173 E(31)[1-4,6,9,12]; A(9)[5,7,9,10,12];
G(60)[1,2]
9 56 E(82)[1,2,6,7,9]; A(18)[1,2,6,12]
5 7 A(43)[1,9,11]; F(57)[1]


E(100)[9]
A(100)[7]
G(100)[1]
A(100)[1]
E(3)[1,3]; A(97)[1,6,9-12]
E(100)[1,2,4,6,7,9,11]
F(100)[2]
E(100)[1-4,6,7,9]
E(38)[1,7,9]; A(12)[6]; G(50)[2]
A(11)[3,6,9]; F(7)[1,2]; G(82)[1,2]


23 64 E(48)[1-7,10]; A(38)[3,6,7,10,11]; F(8)[2,3];
G(6)[1,2]
1 1 G(100)[2]
6 24 E(100)[1,2,7,9]


71 593 E(86)[1-12]; A(13)[2,3,5-12]; F(1)[1,2]


49
54
11

2
49
39
231

79
64
1053
36
1150
19
3
2
212
1003
161
107
32









Table 2-4. Survey Summary (Trichoptera:Spicipalpia).
Coll. Specimens Study Area (% of Total Collections)
Species (n) (n)a [Collection Station Number]


Hydroptilidae
Hydroptila apalachicola Harris et al.
Hydroptila berneri Ross
Hydroptila bribriae Harris
Hydroptila circangula Harris
Hydroptila disgalera Holzenthal & Kelley
Hydroptila eglinensis Harris
Hydroptila hamiltoni Harris
Hydroptila latosa Ross
Hydroptila lloganae Blickle
Hydroptila molsonae Blickle
Hydroptila novicula Blickle & Morse
Hydroptila okaloosa Harris
Hydroptila parastrepha Kelley & Harris
Hydroptila quinola Ross
Hydroptila remita Blickle & Morse
Hydroptila sarahae Harris
Hydroptila waubesiana Betten
Mayatrichia ayama Mosely
Neotrichia armitagei Harris
Neotrichia minutisimella (Chambers)
Neotrichia vibrans Ross
Ochrotrichia apalachicola Harris et al.
Ochrotrichia confusa (Morton)
Orthotrichia aegerfasciella (Chambers)
Orthotrichia baldufi Kingsolver & Ross
Orthotrichia cristata Morton
Orthotrichia curta Kingsolver & Ross
Oxyethira abacatia Denning
Oxyethira chrysocara Harris
Oxyethira elerobi (Blickle)
Oxyethiraflorida Denning
Oxyethira glasa (Ross)
Oxyethira grisea Betten
Oxyethirajanella Denning

Oxyethira kelleyi Harris
Oxyethira lumosa Ross
Oxyethira maya Denning
Oxyethira novasota Ross
Oxyethira pallida (Banks)
Oxyethira pescadori Harris & Keth
Oxyethira savanniensis Kelley & Harris
Oxyethira setosa Denning
Oxyethira verna Ross
Oxyethira zeronia Ross
Rhyacophilidae
Rhyacophila carolina Banksb


3
1
30
3
8
80
25
25+
2
1+
1
8
3
88+
32+
30
135+
13+
8+
7
1
6
1
11+
4
36
2+
14+
1
4+
2+
9+
1
155+

45+
95
27+
59+
9
24
28+
1
1
59+


34 91


A(100)[10]
A(100)[10]
E(100)[1,3,4,11]
E(100)[7,9,12]
E(80)[4,9,11]; A(20)[6]
E(100)[1-5,7]
E(100)[1,4,7]
E(67)[1,3,4,8,9,11,12]; G(33)[1,2]
E(100)[1,4]
E(100)[7]
F(100)[3]
E(100)[1,7]
E(100)[4,10,11]
E(27)[1,3,4,7-9]; A(68)[1-4,6,7,9- 1]; F(5)[1]
E(43)[4,6,7,10,12]; A(57)[7,9-11]
E(100)[1,2,4,6,7,9]
E(80)[2-4,6,7,9-1 ]; F(13)[l]; G(7)[2]
E(46)[4,6,7,9]; A(8)[7]; F(8)[3]; G(38)[1,2]
E(25)[1,9]; G(75)[1,2]
A(100)[7]
A(100)[9]
E(50)[3]; A(50)[7]
A(100)[12]
E(33)[1,7]; A(33)[7,9]; F(23)[l,3]; G(11)[2]
A(50)[7]; F(50)[3]
E(34)[1]; A(33)[2]; F(33)[3]
G(100)[1,2]
E(37)[1,3,7]; A(27)[7,10]; F(9)[1]; G(27)[1,2]
G(100)[2]
E(100)[4,9]
E(50)[1]; G(50)[1]
E(25)[1,6]; A(25)[10]; G(50)[1,2]
A(100)[7]
E(26)[1,3,4,7,9]; A(44)[2,4,6,7,10,11]; F(17)[1-
3]; G(13)[1,2]
E(100)[1,3,4,7-12]
E(37)[l,3,4,7,9]; A(44)[3,7,10,11]; G(19)[1,2]
E(36)[1,3,7,9]; A(64)[1,2,6,7,9-11]
E(6)[4]; A(76)[2,6,7,9-l 1]; F(18)[1,2]
A(75)[10,11]; F(25)[3]
E(86)[1,3,4]; G(14)[[1]
E(67)[1,4,6,9]; G(33)[2]
A(100)[7]
A(100)[7]
E(80)[1,3,4,6,7,9,10]; A(20)[1,2]

E(64)[1-4,7]; A(30)[3,4,7,9-12]; F(6)[1,2]


a + indicates that specimen counts were not made for 1 or more collections.
b Survey area % abundances based on total individuals.









Table 2-5. Survey Summary (Trichoptera:Integripalpia).
Coll. Specimens Study Area (% of Total Specimens)
Species (n) (n) [Collection Station Number]
Beraeidae


Beraea n. sp.
Brachycentridae
Brachycentrus chelatus Ross
Micrasema n. sp.
Micrasema wataga Ross
Calamoceratidae
Anisocentropus pyraloides (Walker)
Heteroplectron americanum (Walker)
Lepidostomatidae
Lepidostoma griseum (Banks)
Lepidostoma latipenne (Banks)
Lepidostoma serratum Flint &
Wiggins
Leptoceridae
Ceraclea cancellata (Betten)
Ceraclea diluta (Hagen)
Ceraclea flava (Banks)
Ceraclea maculata (Banks)
Ceraclea nepha (Ross)
Ceraclea ophioderus (Ross)
Ceraclea protonepha Morse & Ross
Ceraclea resurgens (Walker)
Ceraclea tarsipunctata (Vorhies)
Ceraclea transversa (Hagen)
Leptocerus americanus (Banks)
Nectopsyche candida (Hagen)
Nectopsyche exquisite (Walker)
Nectopsyche paludicola Harris
Nectopsyche pavida (Hagen)
Oecetis cinerascens (Hagen)
Oecetis daytona Ross
Oecetis ditissa Ross
Oecetis georgia Ross

Oecetis inconspicua Complex
Oecetis nocturna Ross
Oecetis osteni Milne

Oecetis persimilis (Banks)
Oecetis sphyra Ross
Triaenodes aba Milne
Triaenodes helo Milne
Triaenodes ignitus (Walker)

Triaenodes n. sp.
Triaenodes ochraceus (Betten &
Mosely)
Triaenodes perna Ross
Triaenodes taenia Ross
Triaenodes tardus Milne


3 4


83
1119
33

540
123

13
13
31


23
35
2
43
17
1
133
13
333
51
22
1
79
832
593
20
3
24
90

875
10
18

47
572
1
4
319

6
1

12
7
6


E(100)[1]


E(98)[2-4,6-12]; A(2)[10]
E(100)[1-12]
A(9)[10]; G(91)[1,2]

E(47)[1-9,11,12]; A(41)[1-3, 5-11]; F(12)[1,2]
E(61)[1-5,7]; A(37)[5,6,8,9,11]; F(2)[1,2]

A(100)[7,9-11]
A(100)[3,5,7,9,11]
E(29)[1,5]; F(71)[1,2]


A(100)[2,4,6,7,11]
E(100)[2,4,6,7,9]
A(100)[1,6]
E(33)[1,4,7,9,10,12]; A(67)[1,2,4,6,10,11]
A(100)[2,4,6]
A(100)[4]
E(1)[6]; A(99)[1,2,4-6,10]
E(100)[2,4,12]
A(100)[1,2,4,6,10]
A(98)[2-6]; F(2)[1]
A(55)[1,6]; F(45)[1,2]
E(100)[4]
E(18)[4,7]; A(82)[1,2,6,7,9,10]
E(100)[1-12]
E(7)[2,7,9]; A(2)[6,10]; G(91)[1,2]
E(15)[1,7,9]; A(80)[1,2,6,11]; F(5)[3]
E(67)[6,7]; G(33)[1]
E(21)[3,4,7]; A(37)[1,7,9,10]; F(42)[1-3]
E(19)[1,3,4,7,9,1 1]; A(17)[2,4,7,9-11]; F(3)[1,2];
G(61)[1,2]
E(69)[1-9]; A(25)[1-11]; F(5)[1-3]; G(1)[1,2]
E(10)[3]; A(80)[1,2,4,7,9,10]; F(10)[2]
E(22)[3,6,7,9]; A(61)[2,6,7,9,11]; F(11)[1,2];
G(6)[2]
E(9)[1,4,6,9]; A(91)[1,2,4,6,7,11,12]
E(62)[2,4-7,9]; A(35)[1,2,4-7,9- 11]; F(3)[1,2,3]
A(100)[10]
A(100)[7,10]
E(6)[1,3,4,6,7,9]; A(45)[1-7,9-11]; F(4)[1-3];
G(45)[1,2]
E(100)[2-4,9]
A(100)[10]

E(100)[2,6,7,9]
A(14)[3]; F(86)[1,2]
A(100)[1,2,6]









Table 2-5. Continued.
Coll. Specimens Study Area (% of Total Specimens)
Species (n) (n) [Collection Station Number]
Limnephilidae
Pycnopsyche antica (Walker) 25 287 E(45)[2-4,6-9]; A(28)[2,3,6,7,9-11]; F(1)[1];
G(26)[1,2]
Pycnopsyche indiana (Ross) 1 1 E(100)[2]
Molannidae
Molanna blenda Sibley 38 184 E(69)[1-5,7]; A(13)[3,7,9-11]; F(18)[1,2]
Molanna tryphena Betten 25 63 E(13)[2,4,6]; A(54)[5-7,10]; F(3)[2,3]; G(30)[1,2]
Molanna ulmerina Navis 7 15 E(100)[1,2,6]
Odontoceridae
Psilotretafrontalis Banks 21 510 E(4)[2,5]; A(17)[3,7-12]; F(79)[1,2]
Phryganeidae
Banksiola concatenata (Walker) 1 1 E(100)[2]
Ptilostomis ocellifera (Walker) 2 2 E(100)[9]
Ptilostomis postica (Walker) 5 5 A(40)[4,7]; F(60)[1,2]
Sericostomatidae
Agarodes crassicornis (Walker) 12 61 E(78)[3,7,9,10]; A(13)[7,10,11]; F(2)[1]; G(7)[1]
Agarodes libalis Ross & Scott 15 248 A(29)[7,8,10,11]; G(71)[1,2]
Agarodes logani Keth & Harris 4 5 F(100)[1,2]
Agarodes ziczac Ross & Scott 23 951 E(100)[1-7,9,12]

Suborder Annulipalpia (Table 2-3)

Family Dipseudopsidae

Three widespread eastern North American Phylocentropus species (P. carolinus, P.

lucidus, P. placidus) were recorded in the survey. Phylocentropus species were most

common within the central panhandle study areas. Phylocentropus lucidus showed the

most restricted distribution and was collected from only stations within the central

panhandle region. Larvae of P. lucidus were found in small, sandy seeps. The other 2

species occurred across a wider geographic area and array of habitat types. However, no

Phylocentropus species were collected from Gold Head Branch even though both P.

carolinus and P. placidus occur in small spring-fed streams of the northern Florida

peninsula.

Family Hydropsychidae

A total of 17 species grouped within 5 genera were inventoried. The genus

containing the greatest number of species was Cheumatopsyche (8), followed by









Hydropsyche (5), Diplectrona (2) and Macrostemum and Potamyia (1 species each).

Species composition and abundances at the sampling stations reflected species stream-

size preferences, as well as large-scale regional differences in the species geographic

ranges.

Genus Cheumatopsyche. Differences in Cheumatopsyche species composition at the

regional and local scales were evident. In the western panhandle region at the Eglin

stations, the 2 most abundant species were Cheumatopsyche gordonae and

Cheumatopsyche petersi, both of which have restricted distributions. Cheumatopsyche

gordonae, widespread and abundant within streams on Eglin Air Force Base, is unknown

outside this region. Accordingly, the species was listed by FCREPA as Threatened.

Cheumatopsychepetersi (listed by FCREPA as Rare) has a somewhat larger range that

includes parts of the western Florida panhandle, coastal Alabama, and Mississippi.

Results of the Eglin collecting showed that C. gordonae occurs most often in the upper-

and middle- stream reaches, while C. petersi is most abundant in larger streams and rivers

such as the Blackwater River. Cheumatopsyche virginica, a Coastal Plain species, was

also common and widespread on Eglin streams but infrequently collected from the

Apalachicola stations. Conversely, Cheumatopsyche edista, also a Coastal Plain species,

was common from the central panhandle stations but was infrequently collected from the

Eglin stations. Cheumatopsyche campyla was collected only from the Apalachicola study

area, mainly from the larger streams or at stations very near the Apalachicola River,

suggesting the river's proximity is an important influence on the occurrence of this

species. Cheumatopsyche pinaca was taken in high numbers from larger streams of the

Apalachicola study area and also at Gold Head Branch, where the highest numbers were








recorded and this was the only Cheumatopsyche species collected. Cheumatopsyche

pettiti, a transcontinental species, collected at most of the central panhandle stations, was

less common on Eglin, perhaps due to abundant populations of C. gordonae and C.

peters.

Genus Diplectrona. Diplectrona modest accounted for the highest total

hydropsychid abundance (1150 specimens). Within the panhandle study areas, this

species was the dominant hydropsychid at steephead and upper ravine-reaches. Larvae

occurred abundantly within leaf packs and small debris-dams. Downstream from ravine-

head reaches this species became less abundant and was replaced by Cheumatopsyche

and Hydropsyche species. Adults appearing to be D. modest were collected from the

steephead at Gold Head Branch, which is east and south of the known range for D.

modest. However, larvae of Diplectrona collected at the steephead all possessed a head

coloration pattern different from typical larvae of D. modest; therefore, this species was

designated Diplectrona sp. A. The coloration pattern of the head is similar to Diplectrona

rossi, which is known only from Schoolhouse Springs in Louisiana. However, the larvae

of Diplectrona sp. A have evenly curved frontoclypeal sutures unlike those ofD. rossi,

which angle sharply near the anterior tentorial pits (Morse & Barr, 1990). Pupal

collections of Diplectrona sp. A may help to clarify the identity of this potential new

species.

Genus Hydropsyche. The Hydropsyche caddisflies inventoried comprise 5 species,

with 3 species (H. elissoma, H. incommoda, and H. rossi) collected in relatively high

numbers from widespread panhandle stations, as compared to the scant collections of H.

betteni and H. decalda. Hydropsyche elissoma was the dominant Hydropsyche species at








the Eglin stations, while the Apalachicola study area supported H. elissoma, as well as

high numbers of H. incommoda and H. rossi. Hydropsyche incommoda was very

abundant in light-trap collections at stations near the Apalachicola River, suggesting that

close proximity to the river is an important influence on the abundance of this species,

which is known to inhabit large Coastal Plain rivers. At Gold Head Branch, the only

Hydropsyche identified were 2 males ofH. decalda. Hydropsyche betteni was collected

from only the FAMU Farm Stream.

Genus Macrostemum. A single species was inventoried, M. carolina. The species is

widespread in the eastern USA and throughout much of northern Florida, but in this

survey was collected solely from Eglin streams, where it was widespread and common in

light-trap collections.

Genus Potamyia. The single North American species of this genus, Potamyiaflava,

was taken in light-trap collections from several stations located near the Apalachicola

River. The species is known to inhabit large rivers, and the adults collected likely moved

from the river into nearby tributary reaches.

Family Philopotamidae

Chimarra species were significant components of the caddisfly fauna at most of the

stream reaches sampled. All five species known to occur in Florida were collected (C.

aterrima, C.falculata, C.florida, C. moselyi, C. obscure). Chimarrafalculata was the

most abundant with nearly 90% of all specimens collected at stations on Eglin, where this

species of restricted distribution appears to have the highest population density. Besides

the Florida panhandle, the geographic range of C.falculata includes the lower Coastal

Plain of Mississippi and Alabama, and parts of southwestern Georgia. Chimarraflorida









has a similar distribution, but extends its range north and east into parts of the Atlantic

Coastal Plain and the Florida peninsula. More than half the C.florida collected came

from Gold Head Branch Station 2. Of the widespread eastern North America species, C.

aterrima and C. obsura, C. aterrima was common and fairly abundant across study areas

at various sampling stations, while C. obscura was collected in low numbers from only a

few central panhandle stations. Chimarra moselyi occurred at many of the panhandle

stations, but usually in low numbers.

Family Polycentropodidae

Polycentropodidae were represented in the survey by 5 genera and 13 species.

Among these are both common and widespread species, as well as 1 disjunct occurrence

of a species never collected before in Florida and 3 restricted-range species that were

listed by FCREPA as Rare or Threatened. Seven species were each represented by fewer

than 10 individuals, and none of the polycentropodids were collected, either as adults or

larvae, in high numbers as were representatives from most of the other annulipalpian

families.

Genus Cernotina. Three species were recorded (C. calcea, C. spicata, C. truncona),

each represented by only 1 individual, indicating that Cernotina species are minor

constituents of the caddisfly fauna of ravine assemblages. Cernotina truncona is an

uncommon Southeastern Coastal Plain species listed by FCREPA as Rare. This species,

collected at Gold Head Branch, is thought to be associated with ponds and lakes and

could have entered the ravine from one of the nearby lakes.








Genus Cyrnellus. The collection of only a single individual of Cyrnellusfraternus

indicates that this widespread and abundant river species is an incidental constituent of

ravine assemblages.

Genus Neureclipsis. The 2 species inventoried (N. crepuscularis, N. melco) were

collected only from Eglin and Apalachicola stations. The widespread and common river

species N. crepuscularis was collected as 1 or 2 individuals from a variety of upper and

lower reach stations, but was collected in greatest abundance at station A12 near the

Apalachicola River. Within the Eglin study area, N. melco, a Southeastern endemic, was

the more common and widespread species.

Genus Nyctiophylax. Three Nyctiophylax species were inventoried (N. affinis, N.

morse, and N. serratus ). Among these 3 species is the widespread North American

species (N. affinis), a southeastern endemic (N. serratus), and N. morse, endemic to

small, cool streams of the western Florida panhandle and coastal Alabama. Within this

survey, N. serratus was the most widespread, occurring across the panhandle stations as

well as from Gold Head Branch. Nyctiopylax morse was collected from a majority of

Eglin stations and overall was the most abundant Nyctiophylax species. Nyctiophylax

morse, along with Polycentropusfloridensis, form a pair of syntopic endemics unique to

the spring-fed streams on Eglin.

Genus Polycentropus. Four Polycentropus species were inventoried (P. blicklei, P.

cinereus, P. clinei, P.floridensis), and all of them have contrasting distributions.

Polycentropus cinereus, a transcontinental species, was the most widespread and

abundant in the survey. Polycentropus blicklei was also widespread, occurring in all

study areas except Eglin. It was collected most frequently and abundantly from the Gold








Head Branch springrun. Other collections of P. blicklei comprised only single

individuals. This species appears to be restricted in Florida to small spring-fed streams.

Also from Gold Head Branch, a single male of Polycentropus clinei was identified. This

species is mainly northeastern in distribution with disjunct populations having also been

found within Minnesota and coastal Alabama (Harris et al., 1991). However, because it

has been sparsely collected on the Coastal Plain, it is difficult to make firm inferences as

to its actual distribution. Polycentropusfloridensis is endemic to spring-fed streams of

the western panhandle and coastal Alabama. It was collected from several different Eglin

streams suggesting that the species is widespread within Eglin, but not as common in

occurrence as other narrow-range endemics such as Nyctiophylax morse,

Cheumatopsyche gordonae, Agarodes ziczac, and Nectopsyche paludicola.

Appropriately, P.floridensis was listed by FCREPA as Threatened.

Family Psychomyiidae

Lype diverse was common at most panhandle stations. In terms of number of

collections (71), it was equaled only by Oecetis inconspicua complex. In northern Florida

habitats other than small spring-fed streams, it is less common.

Suborder Spicipalpia (Table 2-4)

Family Hydroptilidae

Hydroptilidae, also known as microcaddisflies, were the most speciose (44 species

representing 6 genera) family of Trichoptera collected. The genera Hydroptila and

Oxyethira were both represented by 17 species; other genera included: Orthotrichia (4

spp.), Neotrichia (3 spp.) Ochrotrichia (2), and Mayatrichia (1 sp.). Among the

microcaddisflies collected in this study and sent to Dr. Harris for identification were 10








species new to science that have been subsequently described (Hydroptila apalachicola,

H. bribriae, H. eglinensis, H. hamiltoni, H. okaloosa, H. sarahae, H. sykorai,

Ochrotrichia apalachicola, Oxyethira chrysocara, 0. pescadori). Formal descriptions of

these species were provided in Harris et al. (1998), Harris (2002), and Harris and Keth

(2002). As with Integripalpia and Annulipalpia, species composition ofhydroptilids

varied greatly among study areas. Narrow-range endemics were most prevalent within the

Eglin study area. Eglin's cool, clear springruns with abundant vascular plants and

macroalgae provide habitat and food resources that sustain a diverse and unique

assemblage of microcaddisflies. Several of these endemic species are very similar to

more widely occurring Southeastern species (Harris, 2002), and it is likely to a large

degree that the origin of this unique fauna is due to speciation of isolated relict

populations.

Genus Hydroptila. The survey led to the discovery of a number of new species in

this large genus. Of the Hydroptila specimens sent to Dr. Harris for examination, he

discovered 7 species new to science. Five of these (H. bribriae, H. eglinensis, H.

hamiltoni, H. okaloosa, H. sarahae) appear to be endemic to streams on Eglin. The

descriptions for these species, provided in Harris (2002), were based in large part on

specimens collected as a part of this study. The aforementioned species were all collected

from more than one watershed and are probably fairly widespread in the Eglin streams, at

least on the western side of the base where the study area was located. These 5 new

species, because of their restricted distributions, should be considered for special

conservation status. Four additional species (H. circangula, H. lloganae, H. molsonae, H.

parastrepha) with restricted ranges were also collected from the Eglin study area. Of








these, the distributions of H. circangula and H. parastrepha encompass parts of the

western Florida panhandle and coastal Alabama. Hydroptila lloganae (known from TX,

LA, FL, and SC) and H. molsonae, a Gulf Coastal Plain species, were listed by FCREPA

as Rare. Also of note, Hydroptila latosa, a Coastal Plain species, was common within the

Eglin and Gold Head study areas but was not collected from any central panhandle

stations. Endemic to the Apalachicola study area was Hydroptila apalachicola. Described

in Harris et al. (1998), it is known from only 3 specimens collected at Little Sweetwater

Creek (Station A10). It is most similar to H. recurvata, a species endemic to the Black

Warrior system in Alabama. Overall, the most abundant Hydroptila species were 2 very

common and widespread species, H. quinola and H. waubesiana. Other relatively

common and widespread Hydroptila species inventoried included: H. berneri, H.

disgalera, H. novicula, and H. remita.

Genus Mayatrichia. Mayatrichia ayama occurred in all study areas and was most

frequently collected at lower reach stations. This species, widespread in North America,

is a common river species in Florida.

Genus Neotrichia. Three species of Neotrichia (N. armitagei, N. minutisimella, N.

vibrans) were inventoried. Neotrichia minutisimella and N. vibrans were collected from

single stations within the Apalachicola study area. Both are widely distributed over parts

of the eastern and central USA, including most of North and South Florida. Neotrichia

armitagei, on the other hand, is endemic to streams of northern Florida and adjacent

southwestern Georgia. It was described in Harris (1991) from collections made by B.J.

Armitage and M.K. Ward from streams on the western-side of Eglin Air Force Base. In








this study, N. armitagei was also collected from Eglin, but was most frequently collected

from Gold Head Branch where it was abundant at the lower reach (Station G2).

Genus Ochrotrichia. Two Ochrotichia species were recorded in the survey, 0.

apalachicola and 0. confusa, both in low numbers. Ochrotichia confusa was represented

by a single male collected at station A12, a small high-gradient ravine stream located

along the eastern bank of the Apalachicola River. Ochrotrichia apalachicola, a newly-

discovered species, was described in Harris et al. (1998) from a single male collected at

Beaver Dam Creek (Station A7). Subsequently, 5 males were identified from Station E3

of the Eglin study area. These are the only known localities for this species, which

appears to be endemic to the Florida panhandle. Ochrotricha okaloosa was not collected

in the study. This rare species, described in Harris and Armitage (1987), is known only

from the holotype collected 14 August 1985 on Eglin Air Force Base (Turkey Creek at

Base Road 233).

Genus Orthotrichia. Within this small genus, a total of 4 species (0. aegerfasciella,

0. baldufi. 0. cristata, 0. curta) were inventoried. All study areas were represented by at

least 2 species. The widespread and common eastern species, 0. aegerfasciella, occurred

in all study areas and was the most common Orthotrichia collected. Orthotrichia cristata,

also a widespread eastern species, was represented in all panhandle study areas but was

less common (3 collections). Orthotrichia baldufi occurred only within the central

panhandle study areas. This species is northern in distribution, with scattered disjunct

populations within the Southeastern USA. Orthotrichia curta is a primarily southeastern

species and it occurred only within Gold Head study area.








Genus Oxyethira. The 17 species of Oxyethira identified in this survey were equaled

in species richness only by the genus Hydroptila. Narrow-range endemic Oxyethira

species documented in the survey were 0. kelleyi and 0. chrysocara. Oxyethira kelleyi is

widespread and abundant in streams on the western-side of Eglin, but is unknown from

outside this area; the species was described in Harris and Armitage (1987) and was listed

by FCREPA as Threatened. Oxyethira chrysocara, recently described by Harris (2002), is

currently known from only the holotype specimen collected at Gold Head Branch

(Station G2). Additional collecting in and around the type locality is needed to pinpoint

the population center. Three species (0. grisea, 0. setosa, 0. verna) were recorded only

as single individuals from Beaver Dam Creek (Station A7). Oxyethira grisea is unknown

from any other Florida localities, and its occurrence on the Coastal Plain is disjunct.

Oxyethira setosa, listed by FCREPA as Rare, was previously known in Florida from one

other locality, Rocky Creek on Eglin Air Force Base. Another Threatened species, 0.

florida, was recorded from collections made at 2 steephead stations, Eglin (Station El)

and Gold Head (Station G1). Aside from these collections, the species is known in

Florida from only the type locality (Miami) and Temple Terrace near Tampa. Oxyethira

lumosa, a southeastern species, was collected within the Eglin, Apalachicola, and Gold

Head study areas. Dr. Harris closely examined these specimens, and others from northern

Florida and Coastal Plain Alabama, and discovered the presence of 2 closely-related

species, Oxyethira lumosa and a new species, 0. pescadori, described in Harris and Keth

(2002). Subsequently, this new species was also identified from specimens collected by

Dr. Manuel Pescador and others in central South Carolina, suggesting that 0. pescadori

may have a similar distribution to 0. lumosa. Oxyethira elerobi was collected only from









the Eglin study area (Stations E4, E9). The species was listed by FCREPA as Rare, but I

have found additional populations from small blackwater streams in northern Florida,

suggesting that the species is more common than previously thought. Overall, the most

widespread and common Oxyethira species in all study areas was the southeastern

species O.janella. Other Oxyethira species, not already mentioned, collected at 2 or more

study areas included: 0. abacatia, 0. glasa, 0. maya, 0. novasota, 0. pallida, 0.

savanniensis, and 0. zeronia.

Family Rhyacophilidae

Species within the genus Rhyacophila are icons of Appalachian stream biota.

Rhyacophila carolina, unlike other Rhyacophila species, is also found in cool, spring-fed

streams of the Florida panhandle. The species was widespread within the panhandle study

areas and was most abundant at st-3rd order springrun reaches.

Suborder Integripalpia (Table 2-5)

Family Beraeidae

A new species within the genus Beraea was discovered within the steephead at

Station El. Adult males were light trapped on two occasions, and a female was collected

using a beating sheet. The new species will be described in a separate paper. Larvae

remain uncollected, but they probably live in mucky seepage areas within the steephead,

given that Beraea species are noted for their ecological specialization to this type of

microhabitat.

Family Brachycentridae

The Brachycentridae surveyed included Brachycentrus chelatus, Micrasema n. sp.,

and M. wataga. Among these species were an eastern North American species (M.








wataga), a southeastern species (B. chelatus), and a lower Coastal Plain endemic

(Micrasema n. sp.). Brachycentrids were most abundant at lower springrun reaches

having deeper water and submerged aquatic plants. Brachycentrus chelatus was

widespread and common on Eglin, but was collected from only one station (A10) outside

this study area. Micrasema n. sp., described in a dissertation by Chapin (1978), was

widespread and abundant at the lower sampling reaches on Eglin. Larvae were found in

association with submerged macrophytes and macroalgae (Batrachospermum). In

contrast, Micrasema wataga was not collected at the Eglin stations but did occur within

the Apalachicola study area (Station A10) and at Gold Head Branch, where it was most

abundant at the downstream sampling reach (Station G2).

Family Calamoceratidae

The calamoceratid species Anisocentropus pyraloides and Heteroplectron

americanum were collected at panhandle stations, primarily from steephead and upper-

ravine reaches. Their abundance, large size, and feeding habits as detritivores and

shredders of course particulate matter (e.g., leaves), are suggestive of their ecological

importance to ecosystem functioning in these habitats. The more common A. pyraloides

occurred at all such sites; adults were often observed flying during the daytime within the

ravines. Heteroplectron americanum is more restricted in distribution, and in Florida the

species is primarily confined to ravine streams. These isolated Florida populations

probably represent relict populations of a distribution less fragmented during cooler

climate periods, as during the Pleistocene. Likely due to their warm-water intolerance,

neither species has been recorded from peninsular Florida.








Family Lepidostomatidae

Three Lepidostoma species were collected (L. griseum, L. latipenne, L serratum).

The Florida populations are all southern disjuncts and restricted to cool, well-shaded,

spring-fed streams. Lepidostoma griseum and L. latipenne were collected from only the

Apalachicola study area, and both species show Appalachian Highland biogeographic

affininites. Lepidstoma serratum was collected from ravine heads at the FAMU Farm and

on Eglin. The larvae were found on the undersides of chunks of clay-like material that are

most prevalent near spring-source headwall areas. Interestingly, male specimens exhibit

the same genitalic variation noted in Weaver (1988) for specimens collected from

Schoolhouse Springs in Louisiana. The fact that distant populations on the southern

Coastal Plain show the same variation and a disjunct distribution pattern suggests that

these are relict populations, possibly representing an ancestral form of the species that

was forced southward, along with deciduous forest communities, during cooler periods

associated with Pleistocene glacial advances.

Family Leptoceridae

Leptoceridae were the most diverse and abundant integripalpian family collected in

the survey, with leptocerids accounting for nearly 50% of all integripalpian specimens

identified and about 60% of all integripalpian species. Within this family, 32 species

representing 5 genera were inventoried. Species richness for each genus was: Ceraclea

(10), Oecetis (9), Triaenodes (8), Nectopsyche (4), Leptocerus (1). The majority of these

leptocerids are widespread in eastern North America as well as in Florida. However,

there are representatives within Ceraclea, Oecetis, Triaenodes, and Nectopsyche that

either have disjunct populations in Florida or are narrow-range endemics.









Genus Ceraclea. The Ceraclea species recorded have, in general, widespread

distributions over large parts of eastern North America. They occurred most often and in

greatest abundance at the lower-reach stations; at upper-reach ravine stations Ceraclea

species were quite uncommon. Regional differences in species composition of Ceraclea

were evident. The Apalachicola study area supported the most species (8), with C.

protonepha, C. tarsipunctata, and C. transversa being collected primarily from this

region and in the highest overall abundance. Ceracleaflava, C. nepha, and C. ophioderus

were collected only at Apalachicola stations but were less common. Ceraclea maculata

was widespread and common at both Eglin and Apalachicola stations. Ceraclea diluta

and C. resurgens were collected only from Eglin, with C. diluta being the more common.

Harris et al. (1991) reported both of these species from small streams within coastal

Alabama, and from few other Alabama localities. Neither was recorded from streams

surveyed on the eastern part of the reservation (Harris et al., 1982).

Genus Leptocerus. Leptocerus americanus was collected infrequently from a few

central panhandle stations. This species is associated with aquatic macrophytes but was

not collected as larvae or as adults at downstream stations on Eglin where aquatic

macrophytes were abundant. High stream velocities at these stations may account for the

absence since L. americanus is associated more with lentic habitats.

Genus Nectopsyche. Nectopsyche species were common and abundant at middle and

lower reaches. Within steephead- and upper-ravine reaches, Nectopsyche was generally

less abundant. Species composition and abundance varied by region. Three (N. candida,

N. exqusita, and N. pavida) of the 4 species collected range over much of eastern North

America, whereas N. paludicola is endemic to the western Florida panhandle and coastal








Alabama. This species was collected from all Eglin stations, often in great abundance.

Nectopsyche exquisite was the most frequently occurring species from the Apalachicola

study area. Nectopsyche pavida was collected from the Eglin and Apalachicola study

areas, but was most abundant at Gold Head Branch. Collections of this species at the

steephead station (G ) comprised several hundred females but only 4 males, whereas the

collections downstream at station G2 showed a much more equal sex-ratio, suggesting

that females, but not males, were flying upstream. Nectopsyche candida was the least

common species with only a single specimen being identified.

Genus Oecetis. At least nine species of Oecetis were collected, and 4-8 congeners

were typically found at stations where multiple samples were collected. Most widespread

and abundant was the Oecetis inconspicua complex (likely comprising 2 or 3 different

species). Also widespread and very abundant, particularly in lower reaches, was the

southeastern species Oecetis sphyra. Oecetis georgia, another southeastern species, was

collected from all regions but in lower numbers and tended to occur most often in upper

reaches. Oecetis daytona, endemic to the Southeastern Coastal Plain and listed by

FCREPA as Rare, was collected at stations E6, E7, and G1-identified from single

males. The other Oecetis collected (0. cinerascens, 0. ditissa, 0. nocturna, 0. osteni,

and 0. persimilis) are common and widespread eastern species (Floyd, 1995) that were

collected at many of the stations.

Genus Triaenodes. Eight species of Triaenodes were documented in the survey. By

far the most widespread and common was Triaenodes ignitus-collected more often and

in greater total abundance than all other Triaenodes species combined. It occurred at the

majority of stations and appeared to be ubiquitous in streams of all sizes. Larvae were









most frequently found among submerged rootmats of riparian vegetation. Triaenodes

ignitus is widespread and common in streams and rivers across eastern North America.

One species, T. taenia, occurred only in upper ravine-reaches within the Apalachicola

and FAMU Farm study areas. This species was previously thought to be restricted to

mountain streams in the southern Appalachians and spring-fed streams in the Piedmont of

North Carolina and Alabama (Glover, 1996). The cool-modified ravine microclimate is

probably essential for the survival of Florida populations of this Appalachian disjunct.

Four Triaenodes species (T. aba, T. helo, T. ochraceus, T. tardus) were collected only

from the Apalachicola study area. Triaenodes aba and T. tardus are widespread species

typically associated with macrophytes in lentic habitats; the occurrence of single

individuals of these 2 species indicates they do not typically occur in ravine streams. The

southeastern species T. ochraceus and the Coastal Plain species T. helo were collected

from floodplain forest stations (Station A7, A10). Glover (1996) noted that these 2

species have similar habitat preferences, both occurring on cypress roots in small streams.

Triaenodes n. sp. and Triaenodes perna were collected from several Eglin stations.

Triaenodes perna, a close relative of T. helo, ranges over much of eastern North America

and can be found in a variety of lotic and sometimes lentic habitats. Triaenodes n. sp. is

endemic to the Southeastern Coastal Plain, where it is found in small headwater streams.

This species, presently being described by Ken Manuel, was listed in Glover (1996) as

Triaenodes n. sp. A.

Family Limnephilidae

Two species within the genus Pycnopsyche (P. antica, P. indiana) were inventoried.

Pycnopsyche antica was collected from all regions and study areas and generally was








most abundant at middle and lower reach stations. Pycnopsyche species are typically

associated with temperate deciduous forests of eastern North America (Ross, 1963), and

their life history and biology are closely linked with that of deciduous riparian vegetation.

Pycnopsyche species are cool-adapted and in Florida restricted to well-shaded streams

with substantial groundwater inputs, hence the common and abundant occurrence of P.

antica in the streams studied. Pycnopsyche indiana occurred at only 1 station (E2). In

Florida this species has been collected at only a few widely scattered localities

(Rasmussen & Denson, 2000).

Family Molannidae

Molannids occurred commonly along the entire lengths of the streams studied. Three

species were inventoried (M. blenda, M. ulmerina, M. tryphena), all of which are

widespread over much of eastern North America. In this study, clear distinctions were

observed in species biotopes and regional distributions. Molanna blenda was collected in

greatest abundance at ravine-head reaches, while M. tryphena more typically occurred at

lower reach stations. The zonation was only evident in the panhandle where both species

are present. At Gold Head Branch, where it appears that M. blenda is absent, M. tryphena

was more abundant at the steephead reach than at the lower reach on Gold Head Branch.

It appears that M. blenda, which is noted for primarily inhabiting spring-seeps and small

spring-fed streams (Sherberger & Wallace, 1971), competitively excludes M. tryphena

from upper reaches in areas of sympatry. Molanna ulmerina was collected from the

western panhandle at only 3 stations (El, E2, E6). This species also occurs in springruns

of north-central Florida, but was not collected at Gold Head Branch.








Family Odontoceridae

Within Florida, Psilotretafrontalis is largely restricted to the upper reaches of ravine

streams. This species, the only odontocerid known in Florida, was most abundant from

ravine-reach stations within the central panhandle region. Both larvae and adults were

collected in large numbers at the FAMU Farm Stream. Psilotretafrontalis was collected

at only 2 stations on Eglin, and these specimens showed interesting variation in the male

genitalia, specifically in the development of the styles on segment 9. Specimens from

station E5 lacked styles, while specimens from station E2 either lacked styles, possessed

normal styles, or possessed short styles. The lack of styles was used by Denning (1948)

to distinguish P. hansoni from P. frontalis. However, Parker and Wiggins (1987), in their

revision of the genus, considered the specimens they examined lacking styles to be

unusual variants of P.frontalis, and found that in all other respects the specimens were

identical to typical P.frontalis. The variation observed in the Eglin specimens supports

the assertion that this is indeed a variable character, and not a valid basis for considering

them to be separate species. Furthermore, larvae and pupae collected at station E5 were

indistinguishable from those collected at the central panhandle stations.

Family Phryganeidae

Two genera (Banksiola, Ptilostomis) comprising 3 species (B. concatenata, P.

ocellifera, P. postica) were collected in small numbers for this family, whose members,

like Limnephilidae, are primarily northern in distribution. Banksiola concatenata and P.

ocellifera each occurred at only 1 Eglin station, while P. postica was collected on several

occasions from the FAMU Farm stations and at 2 Apalachicola stations.








Family Sericostomatidae

Agarodes species were found to be important components of ravine assemblages

throughout upper and lower reaches. Larvae of species within this genus are noted for

their burrowing habits and preference for spring-fed, sand-bottom streams. Four species

of Agarodes were inventoried, including 2 widespread Southeastern species (A.

crassicornis, A. libalis) and 2 narrow-range endemics (A. logani, A. ziczac). Differences

in species composition were apparent at a regional level. Agarodes crassicornis was the

only species to occur in all study areas. Agarodes libalis was collected from the

steephead springruns in the Apalachicola study area and from the Gold Head Branch

stations. Overall, this species was collected in highest numbers at the Gold Head Branch

steephead. Agarodes ziczac was restricted to the Eglin study area, where it was

widespread and collected in very high numbers. Agarodes ziczac, listed by FCREPA as

Threatened, occurs only in the western Florida panhandle. Results of this survey, and that

of Harris et al. (1982), indicated that the spring-fed headwater streams on Eglin support

the highest populations. The protection of headwater areas on the Eglin Air Force Base is

vital to long-term health of this species. Even more restricted in distribution is Agarodes

logani. This species, described by Keth and Harris (1999), is known only from specimens

collected within the ravine of the FAMU Farm Stream (Stations Fl, F2). The restricted

distribution and apparent small population size suggest that this species is particularly

vulnerable to extinction. Other headwater spring-fed streams around the type locality

need to be sampled in order to better understand its distribution.









Survey Account of Plecoptera

The Plecoptera species inventory, presented in Table 2-6, is discussed below.

Overall, 23 species representing 6 families and 13 genera were identified from among the

116 samples taken at the 29 collecting stations. A total of 759 stonefly specimens were

identified. The family Perlidae contained the lion's share of species (14), followed by

Leuctridae (4), Perlodidae (2), and Nemouridae, Taeniopterygidae, Pteronarcyidae (1

species each). Species richness for the 4 study areas was: Eglin (13 species),

Apalachicola (17 species), FAMU farm (4 species), and Gold Head (2 species). Eight

species found in the Apalachicola study area were not collected at the other study areas,

and 5 species were recorded from only Eglin study area collections. As discussed in the

survey account, the distribution of many stonefly species within and among study areas

appears to be controlled at the local scale by strict habitat requirements, and at a larger

scale by biogeographic affinities. The following account summarizes the survey results.

Family subheadings and species are arranged alphabetically under each of the 2 major

stonefly groupings, Euholognatha and Systellognatha.

Table 2-6. Survey Summary (Plecoptera).
Species Collections Specimens Study Area (% of Total Specimens)
(n) (n) [Collection Station Number]
EUHOLOGNATHA
Leuctridae
Leuctra cottaquilla James 4 9 E(100)[1,5,8,11]
Leuctraferruginea (Walker) 3 3 A(100)[3,7,9]
Leuctra rickeri James 3 5 E(20)[7]; G(80)[1,2]
Leuctra triloba Claassen 2 6 A(33)[11]; F(67)[1]
Neumouridae
Amphinemura sp. 2 4 A(100)[5]
Taeniopterygidae
Taeniopteryx sp. 1 3 A(100)[1]
SYSTELLOGNATHA
Perlidae
Acroneuria abnormis (Newman) 1 1 E(100)[1]
Acroneuria arenosa (Pictet) 13 71 E(7)[4,6]; A(93)[1,2,4-6,9,11]
Acroneuria lycorias (Newman) 40 122 E(26)[4,5,7,9]; A(71)[1-7,9-11]; F(3)[1,2]
Agnetina annulipes (Hagen) 4 9 A(100)[1]





Group Euholognatha (Table 2-6)

Family Leuctridae

Four species of Leuctra were inventoried (L. cottaquilla, L.ferruginea, L. rickeri, L.

triloba). Nymphs of Leuctra were sometimes locally abundant, especially in leaf packs

within upper reaches; unfortunately nymphs could not be identified to species. Therefore,

adult collections from light trapping done during the fall and early spring were used to

determine species composition. The fact that adults of Leuctra are only present in Florida

during the cool months makes them difficult to collect by light, and the low numbers

collected in light-trap samples under-represented their true abundance within ravine

habitats. Of the four species inventoried, L. cottaquilla has the most restricted

distribution. This species is endemic to the western Florida panhandle and parts of

southern Alabama and Mississippi. Within the Eglin study area, it was widespread and

collected at steephead and downstream stations. The other 3 species are widely


\ _I


Table 2-6. Continued.
Species

Eccoptura xanthenes (Newman)
Neoperla carlsoni Stark &
Baumann
Neoperla clymene (Newman)
Paragnetina fumosa (Banks)
Paragnetina kansensis (Banks)
Perlesta placida (Hagen)

Perlesta sp. A
Perlesta sp. B
Perlinella drymo (Newman)
Perlinella zwicki Kondratieff et al.
Perlodidae
Clioperla clio (Newman)
Isoperla dicala Frison
Pteronarcyidae
Pteronarcvs dorsata (Say)


Collections
(n)
19
1

17
6
1
27

1
1
21
1

1
1

9


Specimens
(n)
49
1

311
7
1
67

2
1
55
1

1
9

21


\ IL ~


Study Area (% of Total Specimens)
[Collection Station Number]
A(22)[3,5,8,9,11,12]; F(78)[1]
A(100)[6]

E(0.4)[4]; A(0.6)[1,2]; G(99)[1,2]
E(29)[9]; A(71)[1,2,4,6]
A(100)[9]
E(52)[1,3,4,6-9,1 1]; A(43)[1,2,5,6,9,l 1];
F(5)[1,2]
E(100)[5]
E(100)[6]
E(47)[1-4,6,7,9,12]; A(53)[2,6-11]
E(100)[4]

A(100)[1]
A(100)[2]

E(14)r41: A(86)r1,61









distributed in eastern North America. Leuctra triloba occurred at upper ravine-reach

stations from the Apalachicola and FAMU study areas; these Florida populations are

disjunct from its northern range. Leuctraferruginea also occurred in both of these study

areas, and in the emergence study at the FAMU Farm Stream (see Chapter 4), both

species were found to be syntopic within a 2-m stream section covered by the emergence

trap. Leuctra rickeri was collected from the Eglin and Gold Head study areas, but not

from the central panhandle study areas.

Family Nemouridae

The neumourids were represented by only 2 collections of Amphinemura larvae

(Station A5). It is likely that these specimens belong to Amphinemura nigritta, a species

reported nearby and the only known Amphinemura species in Florida. Amphinemura

species, including A. nigritta, have a prolonged egg diapause (Stewart & Stark, 2002),

which may account for their presence in streams that temporarily go dry. Amphinemura

in Florida appears to be more characteristic of this stream-type than the permanent,

spring-fed systems that were studied.

Family Taeniopterygidae

Taeniopterygidae were represented by only 1 collection (Station Al) comprising 3

nymphs of a Taeniopteryx species. As with the other Euholognathans inventoried,

nymphs usually are not identifiable to species, hence the genus-level determination. The

nymphs likely belong to either T. lita or T. burksi, the only 2 species known to occur

within the central panhandle of Florida.








Group Systellognatha

Family Perlidae

Perlids were, in general, the most common stoneflies collected in the survey, and

undoubtedly they play key ecological roles as predators within the ravine ecosystems

studied. The family was represented by 14 species grouped within 7 genera. Collections

reflected a variety of distributional patterns. Some species inventoried could be

considered ecological generalists (e.g., Perlesta placida, Clioperla clio) while another

species, Eccoptura xanthenes, appears to have strict ecological requirements met only

within the environs of ravine headwaters.

Genus Acroneuria. Three Acroneuria species were inventoried (A. abnormis, A.

arenosa, A lycorias). The widespread North American species, A. abnormis, occurred at

only Station El; in Florida it is more common in rivers than from small spring-fed

streams. Acroneuria lycorias, on the other hand, was common and abundant at panhandle

stations, especially within the Apalachicola study area. The Florida panhandle

populations are disjunct from the species main-range, which encompasses much of the

northern USA and southern Canada east of the Rockies. The absence of this species from

other areas on the Southeastern Coastal Plain suggests that it, like many stonefly species,

is intolerant of high extremes in water temperature, and that panhandle ravine streams are

the key refugia supporting populations on the Southeastern Coastal Plain. Acroneuria

arenosa occurred at various panhandle stations but differs from A. lycorias in that it is

found commonly in stream habitats outside of ravine regions. Although Acroneuria

species do occur within the northern Florida peninsula, none were collected from the









Gold Head study area. The absence of Acroneuria from Gold Head Branch may be due to

the great abundance ofNeoperla clymene.

Genus Agnetina. Nymphs of A. annulipes were collected on several occasions from

Flat Creek (Station Al). This species was absent from steephead and upper ravine-

reaches.

Genus Eccoptura. Closely related to Acroneuria, the monotypic genus is thought to

represent a specialized lineage adapted to spring-fed headwater streams (Stark & Gaufin

1976). The sole member of the genus, E. xanthenes, occurred at steephead and upper-

reach ravine stations within the*Apalachicola and FAMU Farm study areas. It was

particularly abundant within the FAMU Farm ravine. Eccoptura xanthenes is common in

small, springbrooks of the southern Appalachians, and the cluster of populations

occurring in the central panhandle ravines suggests past dispersal between the 2 regions

via Apalachicola/Chattahoochee/Flint watershed connections.

Genus Neoperla. Two species were inventoried (N. carlsoni, N. clymene). Both

occur in a wide array of lotic habitats; within Florida, N. carlsoni is restricted to the

western and central panhandle, while N. clymene is widely distributed over much of

northern Florida. Collections of the 2 species within the panhandle study areas were

scant. In contrast, at Gold Head Branch, N. clymene was very abundant and the only

perlid species to occur.

Genus Paragnetina. Two species were inventoried (P.fumosa, P kansensis).

Nymphs of P.fumosa were collected at several of the larger streams within the

Apalachicola study area, and from Turkey Creek (Station E9) within the Eglin study area.

Paragnetina kansensis was represented by a single female taken from the Apalachicola









study area (Station A9). Neither species appears to be an important component of upper-

reach ravine assemblages.

Genus Perlesta. Nymphs of Perlesta were commonly collected at virtually all

panhandle stations sampled, but unfortunately, nymphs could not be identified to species.

Based on adult specimens, 3 species were inventoried (P. placida, Perlesta sp. A,

Perlesta sp. B). Perlesta placida, which occurs in most Florida panhandle streams and

rivers, was widespread and common from panhandle stations. Perlesta sp. A and Perlesta

sp. B were collected from the Eglin study area (Station E5 and E6, respectively). Perlesta

sp. A was represented by 2 females and Perlesta sp. B by 1 male. The specimens were

sent to Dr. Bill Stark for examination, who concluded that they likely belong to new

species, but that additional specimens are needed for further study. Because 1 collection

comprised females and the other collection was a male specimen, it is possible that the

specimens are of the same species.

Genus Perlinella. Adults of Perlinella were commonly taken in light-trap samples

from the Eglin and Apalachicola study areas. Two species were inventoried (P. drymo

and P. zwicki). Perlinella drymo was common at upper- and lower reach stations within

both Eglin and Apalachicola study areas, while P. zwicki was represented by a single

male specimen taken from Juniper Ck (Station E4). Perlinella zwicki, a Coastal Plain

endemic, more commonly occurs in shifting-sand-bottom rivers such as the Blackwater,

while Perlinella drymo, an eastern species, occurs in a wider array of habitat types,

including small headwater streams.








Family Perlodidae

Only 2 species ofperlodids were collected, Clioperla clio and Isoperla dicala, both

from the Apalachicola study area. A larva of C. clio was collected from Flat Creek

(Station Al), and adults of dicala were taken from Crooked Creek (Station A2).

Clioperla clio occurs across many stream types in northern Florida but apparently is not

an important component of ravine assemblages. Isoperla dicala is rarer within the state

and is known from only a few panhandle streams and rivers.

Family Pteronarcyidae

Pteronarcys dorsata was collected at several lower-reach stations (Stations E4, Al,

A6). Larvae were collected from areas of fast flow where leaf packs accumulated on

snags. This species, widespread in North America, is restricted in Florida to clean,

unpolluted streams and rivers with fast flow. Pteronarcys dorsata did not occur in the

upper reaches of ravine streams.














CHAPTER 3
ANALYSIS OF TRICHOPTERA COMMUNITY STRUCTURE AND
ENVIRONMENTAL RELATIONSHIPS


Based on results of the biodiversity survey of ravine streams across northern Florida

(Chapter 2), it was apparent that there are significant differences in species composition

of the caddisfly and stonefly fauna among the different sampling stations. Examples of

faunal differences between stations and study areas were noted in the survey account, and

species occurrence was generally discussed in terms of species geographic distributions

and habitat associations (e.g., stream size, substrate microhabitats).

From the starting point of understanding the makeup of the stonefly and caddisfly

fauna of the various stations, additional questions can be asked such as: How similar or

different is the fauna between stations? Can the survey results be used to produce a

classification of community types, and if so, what species define a particular community?

What geographic and environmental factors best explain the faunal differences observed

among stations? In an attempt to address these questions, the survey data were analyzed

using multivariate statistical methods. The objectives of the analysis were to i) provide a

classification and description of community types based on faunal similarities between

stations; and ii) investigate key geographic and environmental factors that are predictors

of species composition.








Materials and Methods

Data Set

The faunal survey (see Chapter 2), which resulted in a rather large data set of species

occurrences and abundance values from 29 stations, was used as the starting point in this

analysis. The data set analyzed was limited to caddisfly data, with the stonefly data

excluded from the analyses because of the relatively low species diversity and small

sample-sizes. The low stonefly numbers were in part due to inherent lower diversity and

in part due to the sampling bias that resulted from the heavy reliance on light trapping,

which is ineffective for capturing many stonefly species. The data for microcaddisflies

(Family: Hydroptilidae) were excluded from the analyses because species composition

and abundance data (counts), unlike non-hydroptilid data, were not available for all

blacklight samples. The data set used was further confined to only those stations from

which at least 3 light-trap samples were collected (2 spring/summer samples, and 1 fall

sample). This requirement resulted in 9 stations being dropped from the analyses (E8,

E10-12, Al, A5, A8, A12, F3), leaving the final data set consisting of abundance values

for 72 macrocaddisfly species from 20 stations. Abundance was recorded as the total

number of individuals/species collected at each station. The data set was analyzed using 2

statistical approaches: classification and ordination. Statistical procedures were

performed using the computer software MVSP (Kovach, 1999).

Classification

Stations were classified on the basis of faunal similarity using the clustering

technique Unweighted Pair Group Mathematical Averaging (UPGMA). The purpose of

performing a cluster analysis was to measure similarity of macrocaddisfly faunas among








stations, and to group like faunas into a framework from which community structure and

relationships could be inferred. Similarity among stations was measured as the Spearman

rank order correlation coefficient (rs), which is computed as follows:


6 (R, R2)2
rs 1- k=1
n(n2 -1)

where: R1 and R2 = station pair abundance rank order; k = species abundance; n = total
number of species.

A quantitative measure (i.e., Spearman's Coefficient) was chosen over binary

(presence/absence) similarity measures (e.g., Jaccard) because species varied greatly in

relative abundance among stations; therefore, a presence/absence coding of the data

would not be sensitive to these differences. Spearman rank correlation coefficient was

used instead of other quantitative similarity measures because this procedure uses rank

order of abundance instead of raw totals for abundance. Rank ordering of abundance is

particularly useful in this study because sampling effort (e.g., number of light trap

samples) was not equal among all stations, thus species total abundance values were not

always directly comparable between stations; however, by converting abundance to a

rank order, the relative abundance rankings could be used for faunal composition

comparisons among stations. From the Spearman similarity matrix, the widely used

clustering procedure Unweighted Pair Group Mathematical Averaging (UPGMA) was

used to group stations with like caddisfly faunas into a hierarchy of nested sets. The

results of this analysis, depicted as a dendogram, are used to propose a classification

hypothesis of community structure. Community species composition was characterized

by providing a list of relative abundances for species occurring in each community.









Ordination

An alternative approach to classifying (clustering) station faunas into discrete

groups, which one may infer to represent ecological communities, is to consider that

species composition changes in a more continuous fashion in response to environmental

gradients (e.g., latitude/longitude, altitude, temperature, stream size). Taking this

approach, the stations were ordinated on the basis of their macrocaddisfly faunas to see if

the data were structured in such a way that can be related to environmental gradients

present within the study area. The procedure used, Detrended Correspondence Analysis

(DCA), is widely used with ecological data and has advantages over other ordination

techniques in that both species and station ordinations are produced simultaneously, and

the axes are scaled in standard deviation units that have a definite meaning (Hill &

Gauch, 1980). According to Hill (1979), samples separated by more than 4 standard

deviations will in general have no species in common. Detrended Correspondence

Analysis also is favored over other ordination techniques because it eliminates the "arch"

effect caused by interdependence of axis 1 and axis 2.

Results and Discussion

The results of the data analyses showed that ravine assemblages surveyed from

across northern Florida support a macrocaddisfly fauna that can be classified into distinct

communities. The communities exhibit major differences in species composition that can

be related to geographic region and habitat types. These differences in community

structure attributed to geographic and habitat factors are structured hierarchically. The

large and biogeographically diverse area encompassing the study areas acts as a

geographic template, determining to a large extent the caddisfly fauna of the 3 geographic









regions (i.e., western panhandle, central panhandle, peninsula). The faunas among

regions are sufficiently distinct that stations within regions were more similar to each

other, regardless of habitat factors, than to stations in other regions. Nested within this

geographic template is a habitat template that acts to further shape and define the fauna

into recognizable communities. Discussed below are results of the classification and

ordination analysis as it applies to my interpretation of community structuring.

Community Classification

The similarity matrix (Table 3-1) and UPGMA dendogram (Fig. 3-1) quantify the

degree of similarity in the macrocaddisfly faunal composition among stations. Any 2

stations can be compared by reading the similarity matrix. The dendogram illustrates the

results of the UPGMA analysis. Stations were grouped into a hierarchical framework that

is informative and useful for proposing a community classification. The dendogram

shows that the 2 highly similar (0.77) peninsular stations at the Gold Head Branch

steephead represent a distinct community showing little similarity (0.1) to stations

constituting the panhandle grouping. Likewise, within the panhandle, the Eglin steephead

streams in the western panhandle showed little similarity (0.15) to the central panhandle

station cluster. Within the central panhandle, additional groupings were evident with

large streams within the Apalachicola study area showing only slight similarity (0.27) to

small ravine-headwater streams. The ravine headwater streams, although modestly

similar (0.48), do appear to represent 2 distinct community types, corresponding to

clayhill ravine streams (0.60) and Apalachicola steephead streams (0.62). The Eglin

steephead stream portion of the dendogram suggests that the Eglin stations are not

separable into distinct communities but rather represent a single diverse community.






71


Based on this interpretation of the analysis, 5 distinct communities are recognized


within the data set: Gold Head Branch, Apalachicola Steephead Streams, Clayhill Ravine

Streams, Apalachicola Large Streams, and Eglin Steephead Streams. Community

composition is characterized in terms of what species are present and their relative

abundance (Tables 3-2 and 3-3). These parameters are also used in discussing community

diversity, in that they delimit species richness and eveness. The abundance distribution

for species constituting the different communities is shown in Figure 3-2. Data regarding

the 5 communities are summarized and discussed below.

Table 3-1. Matrix of Spearman rank abundance correlation coefficients for
macrocaddisfly data analyzed in the cluster analysis.
El

El 1 E2
E2 0.54 1 E3
E3 0.65 0.53 1 E4
E4 0.62 0.67 0.71 1 E5
E5 0.60 0.57 0.55 0.59 1 E6
E6 0.51 0.67 0.48 0.68 0.55 1 E7
E7 0.63 0.67 0.68 0.75 0.57 0.68 1 E9
E9 0.40 0.51 0.45 0.53 0.35 0.64 0.65 1 A2
A2 0.04 -0.10 0.03 0.07 0.03 0.15 0.11 0.14 1 A3
A3 0.25 0.26 0.34 0.26 0.30 0.12 0.26 -0.02 0.23 1 A4
A4 -0.05 -0.22 -0.08 -0.04 -0.15 -0.07 -0.08 -0.11 0.61 0.15 1 A6
A6 0.03 -0.09 -0.04 0.02 0.06 0.13 0.08 0.06 0.80 0.20 0.56 1 A7
A7 0.22 0.12 0.41 0.32 0.26 0.20 0.29 0.14 0.42 0.48 0.26 0.33 1 A9
A9 0.34 0.22 0.48 0.40 0.36 0.20 0.31 0.07 0.37 0.54 0.15 0.24 0.66 1 A10
A10 0.15 0.07 0.33 0.24 0.14 0.13 0.24 0.11 0.43 0.36 0.30 0.38 0.76 0.53 1 All
All 0.39 0.15 0.40 0.29 0.36 0.15 0.34 0.14 0.39 0.51 0.30 0.35 0.72 0.66 0.59 1 F1
F1 0.19 0.05 0.35 0.18 0.32 0.02 0.21 0.03 0.31 0.62 0.20 0.24 0.57 0.63 0.40 0.54 1 F2
F2 0.18 0.05 0.29 0.10 0.29 -0.04 0.13 -0.10 0.18 0.58 0.13 0.15 0.49 0.45 0.36 0.40 0.78 1 GI
G1 -0.02 -0.04 0.12 0.01 -0.12 0.05 0.07 0.09 0 0.09 0.02 0.07 0.27 0.08 0.31 0.10 0.09 0.11 1 G2
G2 0.07 0.01 0.15 0.04 -0.10 0.09 0.11 0.15 0.08 0.12 0.04 0.18 0.30 0.16 0.33 0.13 0.12 0.18 0.77 1









Station
G2
SGold Head Branch G1
All
Upper Reaches A9
Apalachicola Steepheads A10
Lower Reaches A7
Ravine Headwater Streams F2
Fl
Clayhill Ravines
Central Panhandle Clayhill RavA3
A4
Apalach. Large Streams A6
Panhandle A2
E9
E5
E6
Eglin Steephead Streams E2
E7
E4
E3
El
I I I I I I i
0.04 0.2 0.36 0.52 0.68 0.84 1
Spearman Coefficient


Figure 3-1. Results of UPGMA cluster analysis of sampling stations using Spearman's
rank order correlation coefficient as the measure of similarity. Analysis was of the
macrocaddisfly data obtained in the survey.

Gold Head Branch

The plot (Fig. 3-2) of species abundance values indicated in Tables 3-2 and 3-3

illustrates that the macrocaddisfly community at the Gold Head Branch springrun is

structured very differently than the other communities. This community is characterized

by low species richness (n=21) and highly uneven abundance distribution, as indicated by

the short and steeply sloping line in Figure 3-2. Despite high-quality habitat of the stream

and presumed wide-breadth of niches, the community is numerically dominated by a

relative few species. In part, this can be explained by the lower caddisfly species richness

of the peninsula versus the panhandle. However, I have collected many other caddisfly

species, and also stonefly species, from small spring-fed streams in the northern









peninsula that I expected to also be present in Gold Head Branch. The low diversity of

the Gold Head Branch community may be due in part to the small size of the watershed

(a short ravine-springrun which ends in a lake); therefore, opportunities for species to

colonize Gold Head Branch are limited. Additionally, land use in the area related to park

development and visitor use, and previous mill operations during the 1930's that included

a dam across the stream, may have caused species extirpations. The Gold Head Branch

community does, however, contain some unique faunal elements, most notable being the

population of Diplectrona sp. A that inhabits the upper reach of the ravine head.

Protection of the steephead from human disturbance is essential to maintain the health of

this species, which is known from only this locality.


100.00



-- Gold Head Branch
10.00 ---Apalach. Steephead Streams

S-*-Clayhill Ravine Streams

SApalach. Large Streams

C ,. H Eglin Steephead Streams
1.00





0.10





0.01 "I I
1 5 9 13 17 21 25 29 33 37 41 45 49 53 57 61
Species sequence

Figure 3-2. Species abundance plots for the 5 macrocaddisfly communities. Species
abundance values according to those listed in Tables 3-2 and 3-3.









Apalachicola steephead streams

The springruns of steephead ravines on the Nature Conservancy Preserve support a

species-rich macrocaddisfly community (54 species) that includes a number of ravine

crenobionts, narrow-range endemics, and disjuncts, as well as widespread habitat

generalists. The numerical dominance of Hydropsyche incommoda reflects the strong

influence of the Apalachicola River on this community. Despite the river's clear faunal

influence, the community's ravine affinity is strong with ravine crenobionts such as

Diplectrona modest, Heteroplectron americanum, Psilotretafrontalis, and Molanna

blenda all occurring in significant numbers. Species that set this community apart include

Lepidostoma griseum, a disjunct species known from no other area of Florida. The

abundance of Hydropsyche elissoma and Chimarrafalculata also distinguishes

Apalachicola steephead streams from clayhill ravine streams. The substantial

contributions of springs to the stream flows, a result of high groundwater storage capacity

of the sandhills, is likely a key factor, as these 2 species are also abundant within Eglin's

steephead springruns.

Clayhill ravine streams

The 3 stations sampled at small clayhill ravine streams in the central panhandle have

a macrocaddisfly community that is somewhat less diverse both in terms of species

richness (38 species) and eveness than the other panhandle communities sampled. Ravine

crenobionts and other caddisflies associated with small spring-fed streams in the Florida

panhandle (e.g., Psilotretafrontalis, Diplectrona modest, Anisocentropus pyraloides,

Molanna blenda, Phylocentropus lucidus, Lepidostoma serratum) are among the most

abundant species. Absent, or far less abundant, are hydropsychids typical of the large









streams and rivers (e.g., Hydropsyche incommoda, Cheumatopsyche pinaca,

Hydropsyche rossi). The community also contains 2 interesting elements: Agarodes

logani, a narrow-range endemic, and Triaenodes taenia, an Appalachian disjunct.

Psilotretafrontalis is especially abundant in this community.

Apalachicola large streams

The large streams of the Apalachicola Bluffs and Ravines Region have a caddisfly

community showing little similarity to headwater ravine communities. The community

has an especially diverse leptocerid component, with 6 of the 10 most abundant species

belonging to this family. In this respect, the Apalachicola large stream community more

generally reflects the warm-adapted fauna of Florida as a whole. The 2 most abundant

species included the large river species, Hydropsyche incommoda, and a habitat

generalist, Cheumatopsyche pinaca. Generally absent are ravine crenobionts and

Appalachian disjuncts, suggesting that habitat and water quality parameters such as

temperature are sufficiently different from ravine headwaters so that the structure of this

community shows little resemblance to that of the small-stream ravine communities of

the central panhandle.

Eglin steephead streams

The caddisfly community of Eglin's steephead ravines and high volume springruns is

distinct and arguably contains the highest concentration of narrow-range endemic

caddisflies anywhere on the Southeastern Coastal Plain. Ravine crenobionts of this

community include species typical of other panhandle ravines such as Diplectrona

modest, Molanna blenda, Heteroplectron americanum, Psilotretafrontalis, and

Lepidostoma serratum. In addition, to this list can be added another ravine crenobiont, a








new species within the genus Beraea. However, what sets the Eglin steephead stream

community most apart from the other communities are the other narrow-range endemics,

3 of which (Micrasema n. sp., Agarodes ziczac, Nectopsychepaludicola) were the most

abundant species within this community. Additional narrow-range endemics such as

Cheumatopsyche gordonae, Cheumatopsyche petersi, Nyctiophylax morse, and

Polycentropusfloridensis are important and unique components of this community,

which are not present in central panhandle or peninsular ravine ecosystems. In total, 20

species of macrocaddisflies occurred only in the Eglin steephead stream community

(Table 3-2)-the other 4 communities each contained only 4 or 5 such species.

The origin and persistence of this unique community appears to be related to both

geographic and habitat factors. The large and insular area encompassing Eglin's

steephead drainage networks contains enough unique stream habitat to act effectively as

an "evolutionary engine" driving speciation. The many narrow-range endemics found on

Eglin appear to have adapted to these rather specialized conditions to such an extent as to

limit their abilities for dispersal and colonization to widespread areas. Means (2000)

hypothesized that past sea-level changes may have acted as an isolating mechanism on

ancestral stocks of Plethodontid salamander populations in steepheads near the coast such

as those on Eglin. Means (2000) suggested that for the steephead ravines on the western

side of Eglin a slight increase in sea level (2-5 meters) would have resulted in the

embayment of the Yellow River, thus causing a saltwater barrier to isolate ancestral

salamander populations within the coastal steephead valleys. It seems plausible that this

isolating mechanism could also be a factor contributing to the area's high degree of

caddisfly endemism.






77



Table 3-2. Macrocaddisfly species composition for Gold Head and Eglin steephead
stream communities, abundance of each species given as percent of total specimens
collected. Note: denotes species recorded from only 1 community.
Gold Head Branch Eglin Steephead Streams Eglin -continued
Species (n=21) (%) Species (n=62) (%) Species (%)
Cheumatopsyche pinaca 38.41 *Micrasema n. sp. 15.41 *Triaenodes n. sp. 0.09
Nectopsyche pavida 26.32 *Agarodes ziczac 13.51 Cheumatopsyche pettiti 0.07
Agarodes libalis 8.47 *Nectopsyche paludicola 9.79 Oecetis ditissa 0.07
Triaenodes ignitus 6.87 Diplectrona modest 9.28 *Beraea n. sp. 0.06
Chimarraflorida 5.03 Oecetis inconspicua Comp. 8.59 Oecetis osteni 0.06
Pycnopsyche antica 3.43 Lype diverse 7.08 Oecetis persimilis 0.06
Chimarra aterrima 3.05 Chimarrafalculata 5.42 Phylocentropus placidus 0.04
Oecetis georgia 2.66 Oecetis sphyra 5.09 Cheumatopsyche edista 0.04
Micrasema wataga 1.45 Anisocentropus pyraloides 3.52 Nyctiophylax serratus 0.04
Polycentropus blicklei 1.06 *Cheumatopsyche gordonae 3.04 Oecetis cinerascens 0.04
*Diplectrona sp. A 0.92 Hydropsyche elissoma 2.01 Hydropsyche incommoda 0.03
Molanna tryphena 0.92 Molanna blenda 1.81 Neureclipsis crepuscularis 0.03
Oecetis inconspicua Comp. 0.53 Pycnopsyche antica 1.65 Oecetis daytona 0.03
Nyctiophylax serratus 0.19 *Macrostemum carolina 1.53 *Ptilostomis ocellifera 0.03
Polycentropus cinereus 0.19 Brachycentus chelatus 1.08 Cheumatopsyche burksi 0.01
Agarodes crassicornis 0.19 Heteroplectron americanum 1.08 Hydropsyche rossi 0.01
*Hydropsyche decalda 0.10 Rhyacophila carolina 0.83 *Cernotina calcea 0.01
*Cernotina truncona 0.05 *Cheumatopsyche petersi 0.78 Ceraclea protonepha 0.01
*Polycentropus clinei 0.05 Chimarraflorida 0.76 *Nectopsyche candida 0.01
Oecetis daytona 0.05 Agarodes crassicornis 0.67 Oecetis nocturna 0.01
Oecetis osteni 0.05 Chimarra moselyi 0.66 *Pycnopsyche indiana 0.01
Nectopsyche pavida 0.57 *Banksiola concatenata 0.01
Chimarra aterrima 0.51
*Ceraclea diluta 0.50
*Nyctiophylax morse 0.48
*Neureclipsis melco 0.43
Polycentropus cinereus 0.43
*Polycentropusfloridensis 0.34
Cheumatopsyche virginica 0.31
Psilotretafrontalis 0.30
Triaenodes ignitus 0.29
Oecetis georgia 0.23
*Molanna ulmerina 0.21
Nectopsyche exquisite 0.20
Ceraclea maculata 0.17
*Triaenodes perna 0.17
Lepidostoma serratum 0.13
*Ceraclea resurgens 0.13
Molanna tryphena 0.11
Phylocentropus carolinus 0.10










Table 3-3. Macrocaddisfly species composition for central panhandle communities,
abundance of each species given as percent of total specimens collected. Note: denotes
species recorded from only 1 community.


Apalach. Steephead Streams Clayhill Ravine Streams Apalach. Large Streams


Species (n=54) (%) Species (n=38)
Hydropsyche incommoda 26.6 Psilotretafrontalis
Diplectrona modest 11.3 Diplectrona modest
Anisocentropus pyraloides 7.6 Anisocentropus pyraloides
Oecetis inconspicua Comp. 7.2is i a
Hyr y r 5.8 Oecetis inconspicua Comp.
Hydropsyche rossi 5.8
Agarodes libalis 3.8 Molanna blend
Pycnopsyche antica 3.5 Phylocentropus lucidus
Hydropsyche elissoma 2.9 Cheumatopsyche pettiti
Psilotreta frontalis 2.8 Lype diverse
Chimarra falculata 2.5 Lepidostoma serratum
Chimarra aterrima 2.5 Triaenodes ignitus
Oecetis sphyra 2.0 Oecetis sphyra
Triaenodes ignitus 1.9
Nectopsyche exquisite 1.8 Rhyacophila carolina
Molanna tryphena 1.7 Leptocerus americanus
Heteroplectron americanum 1.4 Oecetis ditissa
Lype diverse 1.2 Lepidostoma latipenne
Molanna blenda 1.1 *Triaenodes taenia
Cheumatopsyche pinaca 1.1 Cheumatopsyche pinaca
Cheumatopsyche pettiti 1.0 Chimarra aterrima
Rhyacophila carolina 1.0
Phylocentropus lucidus 0.9 Polycentropus cinereus
Polycentropus cinereus 0.9 Cheumatopsyche edista
Cheumatopsyche edista 0.9 Pycnopsyche antica
Phylocentropus carolinus 0.7 *Agarodes logani
*Lepidostoma griseum 0.7 Chimarra obscura
Chimarra florida 0.6 Ceraclea transversa
Oecetis georgia 0.6 Phylocentropus carolinus
Oecetis ditissa 0.4
Agarodes crassicornis 0.4 *Nyctiophylax affinis
Ceraclea maculata 0.3 Polycentropus blicklei
Oecetis persimilis 0.3 Oecetis georgia
Potamyia flava 0.2 Ptilostomis postica
Neureclipsis crepuscularis 0.2 *Hydropsyche betteni
Lepidostoma latipenne 0.2 Heteroplectron americanum
Oecetis nocturna 0.2 Oecetis osteni
*Triaenodes helo 0.2
Cheumatopsyche virginica 0.2 Cheumatopsyche burksi
Micrasema wataga 0.2 Hydropsyche elissoma
Ceraclea cancellata 0.2 Hydropsyche incommoda
Oecetis osteni 0.2 Oecetis nocturna
Cheumatopsyche campyla 0.1 Molanna tryphena
Chimarra obscura 0.1 Agarodes crassicornis
Brachycentus chelatus 0.1
Ceraclea tarsipunctata 0.1
Phylocentropus placidus 0.1
*Cernotina spicata 0.1
Polycentropus blicklei 0.1
Ceraclea protonepha 0.1
Nectopsyche pavida 0.1
Oecetis cinerascens 0.1
*Triaenodes aba 0.1
* Triaenodes ochraceus 0.1
Ptilostomis postica 0.1


(%) Species (n=44) (%)
39.5 Hydropsyche incommoda 23.9
22.2 Cheumatopsychepinaca 13.8
7.3 Oecetis sphyra 9.2
3.4 Ceraclea tarsipunctata 9.2
3.4 Ceraclea protonepha 7.9
3.2 Triaenodes ignitus 5.4
3.2 Oecetis inconspicua Comp. 3.4
2.4 Ceraclea transversa 3.1
2.1 Hydropsyche rossi 2.7
1.4 Anisocentropuspyraloides 2.3
1.3 Oecetis persimilis 2.1
1.1 Phylocentropus carolinus 1.6
0.9 Nectopsyche exquisite 1.5
0.8 Ceraclea cancellata 1.3
0.7 Potamyiaflava 1.2
0.7 Ceraclea maculata 1.2
0.6 Lype divers 1.1
0.6 *Ceraclea nepha 1.1
0.6 Cheumatopsyche campyla 1.1
0.5 Cheumatopsyche edista 0.6
0.5 Leptocerus americanus 0.6
0.5 Nectopsychepavida 0.5
0.4 Oecetis cinerascens 0.5
0.4 Oecetis osteni 0.5
0.3 Pycnopsyche antica 0.5
0.3 Phylocentropus placidus 0.4
0.3 Cheumatopsyche pettiti 0.4
0.3 Polycentropus cinereus 0.3
0.3 Heteroplectron americanum 0.3
0.2 Hydropsyche elissoma 0.2
0.2 Chimarra moselyi 0.2
0.2 Oecetis georgia 0.2
0.1 Oecetis nocturna 0.2
0.1 Triaenodes tardus 0.2
0.1 Diplectrona modest 0.1
0.1 Molanna tryphena 0.1
0.1 Phylocentropus lucidus 0.1
0.1 Neureclipsis crepuscularis 0.1
Nyctiophylax serratus 0.1
Polycentropus blicklei 0.1
*Ceracleaflava 0.1
*Ceraclea ophioderus 0.1
Ptilostomis postica 0.1
Rhyacophila carolina 0.1






79


Ordination Analysis

The results of the detrended correspondence analysis used to ordinate sampling

stations are presented in Figures 3-3 to 3-5. The first ordination (Fig. 3-3) is for all 20

stations. This analysis resulted in stations being ordinated along 5 axes with eigenvalues

of 0.65, 0.35, 0.14, 0.05, and 0.03, respectively. The stations as plotted along the first 2

axes shown in Figure 3-3 account for about 35% of the variation in the data. Because of

the presence of additional axes beyond axis 1 and 2, interpretation of the graph is rather

tenuous. However, it does appear that in general axis 1 is related to a geographic gradient

(longitude), and axis 2 is related to a stream-size gradient. This assessment is in general

agreement with the cluster analysis, wherein basal groupings corresponded with station

location within a particular region (western panhandle, eastern panhandle, and peninsula).



3.9--

F1 F2
3.1-

A3
El E5 A All
A A A9 A10
2.3-- E3 A A4
u X A A A7
C) < A G1 G2
S < A6 A A
on Q
0 01.5E E7
U AA A2
E4

0.8- A


E9
o0.0o I
0.0 0.8 1.5 2.3 3.1 3.9
western panhandle DCA Axis 1 peninsula

Figure 3-3. Scatter plot of DCA ordination of 20 stations across northern
Florida. Analysis based on macrocaddisfly survey data.








The wide spread of the central panhandle stations along axis 1, however, indicates that

longitude alone does not explain station placement along axis 1.

In order to investigate gradients related to habitat variables more closely, the western

panhandle and central panhandle stations were separately ordinated to remove the effect

region imparts on the ordination. Both ordinations (Figs. 3-4 and 3-5) indicated that

variation between stations is largely explained in terms of a stream-size gradient. For the

ordination of Eglin stations (Fig. 3-4), axis 1 explained about 43% of the variation in

station scores. In this graph, upper reach stations are placed at one end axis 1 and Station

E9, the station with greatest discharge, at the other end; stations from middle reaches

were clustered about midway along axis 1. A similar result was obtained for the central

panhandle stations (Fig. 3-5). In this ordination, axis 1 explained about 38% of the

variation in station scores. Large stream stations were placed at one end of the axis and

ravine headwater reaches at the other end.

Results of both cluster analysis and ordination suggest that caddisfly community

composition is controlled both by geographic and habitat factors. The study areas, which

span much of northern Florida, are distant enough from one another that the geographic

ranges of many species do not include all study areas. The diverse array of component

species geographic distributions acts as a base template upon which community

composition is structured. From this template, species composition at a given site is then

fine-tuned by habitat factors related to stream size and ravine type.
















2.2+


steephead reaches


1.6



U
o 1.1-



o.5-


middle reaches


lower reaches


E4
A E6
A


E2
A
).5 1.1
DCA Axis 1


Figure 3-4. Scatter plot of DCA ordination of steephead stream stations on
Eglin Air Force Base in the western panhandle. Analysis based on
macrocaddisfly survey data.


large streams


ravine springruns


A9 A3
A A


F2 F1
A A


1
1.3
DCA Axis 1


Figure 3-5. Scatter plot of DCA ordination of central panhandle stations.
Analysis based on macrocaddisfly data.


0.0 L-
0.0


0.6--


#2
-
0.0
0.0


L


0


2.7-r-


3.2--














CHAPTER 4
TRICHOPTERA AND PLECOPTERA FLIGHT SEASONALITY, AND
ADULT EMERGENCE IN A RAVINE SPRINGRUN

Information regarding adult emergence and flight seasonality is of key importance to

the study of caddisfly and stonefly life histories. The adult stage is a relatively short-lived

life stage when caddisflies and stoneflies must emerge, find mates, and oviposit.

Additionally, it is during the adult stage when dispersal to new habitats is greatest.

Despite the biological importance of the adult stage, relatively little is known of the

biology of adults as compared to larvae. Because the survey resulted in large numbers of

adult collections, these data can provide important baseline data on flight seasonality for

the species and populations that were sampled. Towards this goal, collection dates were

examined and flight seasonality was characterized for various species recorded in the

survey. Additionally, species diversity and emergence phenology of caddisfies and

stoneflies inhabiting the ravine springrun at the FAMU Farm study area were investigated

using emergence traps.

Materials and Methods

Observed Flight Seasons

Flight seasonality was characterized for caddisfly and stonefly species inventoried as

a part of the biodiversity survey of ravine ecosystems in northern Florida (Chapter 2). A

description of ravine habitats and the stream systems surveyed for caddisflies and

stoneflies is presented in Chapter 1, included are maps (Figs. 1-1 to 1-5) showing the

locations of the 4 study areas and 29 sampling stations. The monthly occurrence of adult








caddisfly and stonefly species was obtained from the adult collection data obtained in the

survey. Data on monthly occurrences for each species from all 29 sampling stations were

combined and tabulated in order to give a general picture of adult flight-seasonality for

each species within the study area as a whole. Adult specimens were captured primarily

by light trapping and mainly during spring and fall. No light trapping was done during

January, February, and July. The number of light-trap samples collected in each month is

as follows: March (20), April (26), May (14), June (22), August (4), September (3),

October (15), November (8), December (4).

Emergence Study

The emergence study was conducted at the FAMU Farm study area (Fig. 1-4) within

a ravine located on the property of the Florida A&M University Research and Extension

Center in Gadsden County (see Chapter 1

for a description of the study area). Adult

aquatic insects emerging from the ravine

springrun at the FAMU Farm were

sampled using emergence traps. The

aquatic insect emergence trap used

(BioQuip Item No. 2829) (Fig. 4-1)

consisted of a 2.4-m-high tent structure

suspended over a 4 m2 area. The front and

back of the trap is cut 0.4 m shorter than

the sides so that water is able to flow
Figure 4-1. Emergence Trap 1 installed
under without obstruction. The traps were over the FAMU farm springrun, near the
ravine head.








each suspended over the stream from parachute cord tied between 2 trees. The sides and

covers were staked so the bottom flaps were in contact with the stream bank on the sides

and in contact with the stream surface in the front and back, thereby preventing emergent

insects from escaping. Aquatic larvae were not confined and could move into or out from

under the traps. The tent material consisted of Lumite screen with 32 x 32 mesh (530

micron openings). Access to the inside of the trap was through a 1.2 m slit-opening

sealed with Velcro.

Two traps were deployed over the FAMU Farm ravine springrun. Trap 1 was located

about 20 meters below the base of the ravine head, and Trap 2 was placed about 70

meters below the ravine head. The composition of benthic substrate within Trap 1 was

estimated to consist of sand (45%), gravel (20%), mud/silt (15%), snags/leaf packs (15%)

and roots (5%). Substrate within Trap 2 comprised sand (65%), roots (15%), mud/silt

(10%), gravel (5%) and snags/leaf packs (5%). Trap 1 was installed 12 March 1998 and

samples collected until May 24th after which time the cord used to suspend the trap broke

due to an unknown reason, causing the trap to collapse. The trap was then reinstalled on

25 June 1998, and sampling continued for a full year until 5 July 1999. Trap 2 was

installed 31 March 1999 and deployed until 5 July 1999.

Insect samples from each trap were collected at about 10-day intervals, resulting in

41 samples from Trap 1 and 8 samples from Trap 2. Stonefly and caddisfly adults trapped

within the tent were collected by aspiration for smaller specimens and by using a soft-

touch forceps for capturing larger specimens. All specimens were placed in 80% ethyl

alcohol. Usually the insects were found resting either at the peak or on the sides and

covers of the trap. Sufficient time (usually about 25 minutes) was spent searching and









collecting within the tent until no additional specimens could be found. Significant

numbers of emergent insects were likely missed in sampling as a result of mortality

during sampling intervals due to spider predation and other mortality factors.

Specimen identification and data analysis. Trichoptera and Plecoptera specimens

collected from the emergence traps were identified to species in most cases. The

microcaddisflies (Family:Hydroptilidae) were identified by Dr. Steven Harris, and I

identified all other specimens. Synoptic voucher collections will be deposited at the

following collections: Florida A&M University Aquatic Insect Collection, Clemson

University Arthropod Collection, and the personal collection of the author. The

deposition of type material for Hydroptila sykorai is given in Harris (2002). For each

sample, the number of males and females of each species were recorded. Collection data

were tabulated, and emergence phenology for the most abundant species was graphed.

Results and Discussion

Observed Flight Seasons (Trichoptera)

In warm temperate climates such as found in northern Florida, adult caddisflies are

potentially present at any time of year, and as expected, specimens were collected during

all months when light trapping was conducted. Based on the monthly occurrence data

from the survey, species could be grouped into 3 flight-season categories: spring/summer

species (S), fall species (F), and those species with extended flight seasons (E) (Table 4-

1). Species with fewer than 4 occurrences were not classified as to their flight seasonality.

The highest number of species fell into the extended fight season category (68 spp.),

followed by spring/summer species (28 spp.) and fall species (1 sp.).

Spring/summer species. The 28 species in this group were collected only during the









spring and summer months (March-September). Some species within this grouping

appeared to be early spring species such as Ceraclea resurgens, while most species

occurred as adults throughout spring (March-May) and into the summer period (June-

September). The low number of samples collected during July, August, and September

make it difficult to determine the extent of species still present as adults during the late-

summer period.

Because sampling intensity was greatest in the spring, results from this period were

examined separately for patterns and trends in species occurrences. During the spring

light-trapping period, caddisfly species richness was highest in May with a total of 105

species being collected, April samples accounted for 96 species, followed by June (81

spp.) and March (67 spp.). Some differences were noted between major groups of

caddisflies in terms of changes in species richness throughout the spring months (Fig. 4-

2). Relatively few leptocerid species were present in March, followed by a sharp increase

in leptocerid species richness to a peak in April, and then a gradual decrease in May and


40
35





Leptoceridae


Integripalpia (other than Leptoceridae)
--I-ntegripalpia (other than Leptoceridae)
0 ----------------------
March April May June
Month of Collection

Figure 4-2. Caddisfly species richness recorded from light trap
samples collected during spring months. Number of samples (n) for
each month as follows: March (20), April (26), May (14), June (22).









June samples. Hydroptilid species richness increased steadily to a peak in May, followed

by a sharp decrease in richness in the June samples. Integripalpia (other than

Leptoceridae) were present at fairly constant levels of species richness throughout the

spring, while the annulipalpians showed a gradual increase to a peak in May followed by

a gradual drop off in species richness in June samples. Most of the cool-adapted species

fall into these last 2 groups, so it is not surprising that a relatively high percentage of the

total species collected in these groups were present in the early spring when air and water

temperatures tend to be lower. Conversely, warm-adapted species prevalent within

Leptoceridae and Hyroptilidae should have a lower percent occurrence in the early spring

and increased occurrences later in the spring.

Fall species. The fall species constituted the smallest seasonality grouping. The most

conspicuous caddisfly within fall light-trap samples was Pycnopsyche antica. In all

panhandle populations sampled, adults of this species were collected from October to

December; however, members of the peninsular population at Gold Head Branch were

also collected in March, May, and June, suggesting that the life history of this population

differs in some significant way from that of the panhandle populations. Aside from

panhandle populations of Pycnopsyche antica, the only other species restricted to the fall

season was Lepidostoma griseum (collected only during October). Weaver (1988)

reported flight dates from 20 June 27 September for L. griseum across its northern

range, suggesting that the Apalachicola ravines population may emerge later in the year

than do northern populations.

Extended flight-season species. The greatest number of caddisfly species (68)

occurred as adults during both spring/summer and fall seasons. A number of underlying









factors may account for species exhibiting flight periods extending over a wide monthly

range. These include: bivoltine life histories, univoltine populations with separate spring

and fall cohorts, continuous generations showing no distinct cohort structure, and

different emergence phenologies between conspecific populations. An example of

differing emergence phenologies between populations appears to be the case with

Cheumatopsyche virginica, which was collected from March to June on Eglin, but was

collected during March, August, and October from the Apalachicola study area.

Psilotretafrontalis also presents a good example. This species appeared to have a narrow

period of emergence (April, May) in most populations sampled, but within some streams

in the Apalachicola study area adults also occurred in October, indicating part of the

population emerges in the fall.

Table 4-1. Monthly occurrences of adult Trichoptera collected in survey.
>. Monthly Occurrence of Adults Collected by Light Trapping
(Number of Light Tra Samples)
->

Species Z o r -


ANNULIPALPIA
Dipseudopsidae
Phylocentropus carolinus 22 X X X X X X X X E
Phylocentropus lucidus 21 X X X X X X X X E
Phylocentropus placidus 9 X X X X X X E
Hydropsychidae
Cheumatopsyche burksi 2 X X ---
Cheumatopsyche campyla 7 X X X S
Cheumatopsyche edista 15 X X X X S
Cheumatopyche gordonae 21 X X X X X X E
Cheumatopsyche peters 16 X X X X S
Cheumatopsyche pettiti 25 X X X X X X X E
Cheumatopsyche pinaca 30 X X X X X X X E
Cheumatopsyche virginica 19 X X X X X X E
Diplectrona modest 58 X X X X X X X X E
Diplectrona sp. A 3 X X X E
Hydropsyche betteni 2 X X ---
Hydropsyche decalda 1 X ---
Hydropsyche elissoma 37 X X X X X X S










Table 4-1. Continued.
n > Monthly Occurrence of Adults Collected by Light Trapping
_(Number of Light Trap Samples) -

S> M > c o M 0 M
Species r- C < ^ < gD
o) 0 S~0 w ~ 00
---- ---- ---- -- -- -- -
Hydropsyche incommoda 35 X X X X X X X E
Hydropsyche rossi 23 X X X X X X E
Macrostemum carolina 10 X X S
Potamyia flava 8 X X X X X E
Philopotamidae
Chimarra aterrima 27 X X X X X X E
Chimarrafalculata 47 X X X X X X X E
Chimarraflorida 26 X X X X X E
Chimarra moselyi 9 X X X S
Chimarra obscura 5 X X X X X E
Polycentropodidae
Cernotina calcea 1 X ---
Cernotina spicata 1 X
Cernotina truncona 1 X ---
Cyrnellus fraternus 1 X ---
Neureclipsis crepuscularis 10 X X X X X E
Neureclipsis melco 13 X X X X X E
Nyctiophylax affinis 1 X ---
Nyctiophylax morse 10 X X X S
Nyctiophylax serratus 6 X X X E
Polycentropus blicklei 12 X X X X X X E
Polycentropus cinereus 23 X X X X X X E
Polycentropus clinei 1 X ---
Polycentropusfloridensis 6 X X X E
Psychomyiidae
Lype diverse 71 X X X X X X X X E
SPICIPALPIA
Hydroptilidae
Hydroptila apalachicola 1 X ---
Hydroptila berneri 1 X ---
Hydroptila bribriae 7 X X X X E
Hydroptila circangula 3 X X ---
Hydroptila disgalera 5 X X X S
Hydroptila eglinensis 9 X X X E
Hydroptila hamiltoni 4 X X S
Hydroptila latosa 15 X X X X X E
Hydroptila lloganae 2 X ---
Hydroptila molsonae 1 X ---
Hydroptila novicula 1 X ---
Hydroptila okaloosa 3 X X ---
Hydroptila parastrepha 3 X X ---
Hydroptila quinola 22 X X X X X E
Hydroptila remita 14 X X X X X X E
Hydroptila sarahae 9 X X X E










Table 4-1. Continued.
S> Monthly Occurrence of Adults Collected by Light Trapping
_(Number of Light Trap Samples 3


Species Z c h ) < X
2'~-' -s 0
00


Hydroptila waubesiana 15 X X X X X X X E
Mayatrichia ayama 13 X X X X X X X E
Neotrichia armitagei 8 X X X X E
Neotrichia minutisimella 1 X --
Neotrichia vibrans 1 X ---
Ochrotrichia apalachicola 2 X
Ochrotrichia confusa 1 X ---
Orthotrichia aeerfasciella 9 X X X X X X E
Orthotrichia balduf 2 X ---
Orthotrichia cristata 3 X X ---
Orthotrichia curta 2 X ---
Oxyethira abacatia 11 X X X S
Oxyethira chrysocara 1 X ---
Oxyethira elerobi 2 X ---
Oxyethira florida 2 X --
Oxyethira glasa 8 _X X X X X E
Oxyethira grisea 1 X ---
Oxyethirajanella 23 X _X X X X X X X E
Oxyethira kelleyi 13X X X X X E
Oxyethira lumosa 16 X X X X X E
Oxyethira maya 14 X X X X X X E
Oxyethira novasota 17 X X X X X X X E
Oxyethira pallida 4 X X X E
Oxyethira pescadori 7 X X X S
Oxyethira savanniensis 6 X X X __ X X E
Oxyethira setosa 1 X ---
Oxyethira verna 1 X -
Oxyethira zeronia 10 X X X X X X E_
Rhyacophilidae
Rhyacophila carolina 34 X x X X X X X E
INTEGRIPALPIA
Beraeidae
Beraea n. sp.3 X -
Brachycentridae
Brachycentrus chelatus 21 X X X X X E
Micrasema n. sp. 27 X X X X X X E
Micrasema wataga 9 X X X X X E
Calamoceratidae
Anisocentropus pyraloides 49 X X X XX X X X E
Heteroplectron americanum 19 X X X X X X E
Lepidostomatidae
Lepidostoma griseum 4 X F
Lepidostoma latipenne 9 X X X X X X E
Lepidostoma serratum 7 X X X X X E










Table 4-1. Continued.
> Monthly Occurrence of Adults Collected by Light Trapping
S(Number of Light Trap Samples






Leptoceridae
Ceraclea cancellata 8 X X X X S
Ceraclea diluta 8 _X X S
Ceraclea flava 2 X X ---
Ceraclea maculata 19 X X X X S
Ceraclea nepha 5 X S
Ceraclea ophioderus X ---
Ceraclea protonepha 10 X S
Ceraclea resurgens 4 X S
Ceraclea tarsipunctata 9 X S
Ceraclea transversa 11 X X X S
Leptocerus americanus 5 X X S
Nectopsyche candida 1 X ---
Nectopsyche exquisite 15 X X X X S
Nectopsyche paludicola 40 X X X X X X E
Nectopsyche pavida 19 X X X X X E
Oecetis cinerascens 10 X X X X X S
Oecetis daytona 3 X X X E
Oecetis ditissa 16 X X X X X X E
Oecetis georgia 27 X X X X X X E
Oecetis inconspicua Complex 71 X X X X X X X X E
Oecetis nocturna 9 X X X X X E
Oecetis osteni 15 X X X X X E
Oecetis persimilis 17 X X X X X E
Oecetis sphyra 26 X X X X X S
Triaenodes aba 1 X ---
Triaenodes helo 2 X --
Triaenodes ignitus 50 X X X X X X X X E
Triaenodes n. sp. 6 X X X S
Triaenodes ochraceus 1 X --
Triaenodes perna 5 X X X S
Triaenodes taenia 4 X X X S
Triaenodes tardus 4 X X S
Limnephilidae
Pycnopsyche antica 25 X X X X X X E
Pycnopsyche indiana 1 X ---
Molannidae
Molanna blenda 38 X X X X X X X X E
Molanna tryphena 25 X X X X X X X E
Molanna ulmerina 7 X X X X E
Odontoceridae
Psilotretafrontalis 21 X X X X X E
Phryganeidae
Banksiola concatenata 1 X









Table 4-1. Continued.
n > Monthly Occurrence of Adults Collected by Light Trapping
C- (Number of Light Trap Samples) _

> > co
Species Z O


Ptilostomis ocellifera 2 __ X X --
Ptilostomis postica 5 X X X E
Sericostomatidae
Agarodes crassicornis 12 X X X X_ S
Agarodes libalis 15 X X X X X E
Agarodes logani 4 X X X X E
Agarodes ziczac 23 _X X X X X E

Observed Flight Seasons (Plecoptera)

The monthly occurrences of stoneflies collected as a part of the biodiversity survey

(Table 4-2) revealed several distinct adult seasonality patterns. The Leuctra species,

members of the euholognathan grouping-what are traditionally called the "winter

stoneflies"-occurred as adults only during the coolest months of the year (October-

March). It is interesting to note that not all Leuctra species typify the "winter stonefly"

designation in northern latitudes where a number of species typically emerge during

spring and summer. In populations of Leuctra spp. throughout northern Florida, however,

emergence is only from fall to early spring.

The Systellognatha consisted of mainly spring/summer species. The only

systellognathans collected in the fall were Neoperla clymene and Perlesta placida.

Neoperla clymene, the dominant stonefly at Gold Head Branch, occurred in spring,

summer, and fall samples but was most abundant in May and June samples. Perlesta

placida mainly occurred as adults in spring and summer, although there was one

anomalous record of a single individual collected in November. The other

systellognathans showed a narrower seasonal-range in adult occurrence. Perlinella drymo









appeared rather early in the spring (March and April) and did not persist long, suggesting

that growth and development are more tightly synchronized in this species than that of

most stonefly species. Eccoptura xanthenes was collected as adults only in late-May and

June. The common perlid, Acroneuria arenosa was most abundant in June, while

Acroneuria lycorias adults were more prevalent in April and May. Other stoneflies

collected in the survey, but not mentioned here, were either collected only as nymphs or

were collected as adults too infrequently to make generalizations as to their adult flight

seasons. Adult collection periods for all species of stoneflies known to occur in Florida

were summarized in Rasmussen et al. (2003).

Table 4-2. Monthly occurrence of adult Plecoptera recorded in survey.
0 > Monthly Occurrence of Adults Collected by Light Trapping
o g (No. of Light trap Samples)
> H

Species Z 2: 0 r o < C


EUHOLOGNATHA
Leuctridae
Leuctra cottaquilla James 4 X X X
Leuctraferruginea (Walker) 3 X X
Leuctra rickeri James 1 X
Leuctra triloba Claassen 2 X X
Neumouridae
Amphinemura sp. 0
Taeniopterygidae
Taeniopteryx sp. 0
SYSTELLOGNATHA
Perlidae
Acroneuria abnormis (Newman) 1 X
Acroneuria arenosa (Pictet) 11 X X
Acroneuria lycorias (Newman) 5 X X X
Agnetina annulipes (Hagen) 0
Eccoptura xanthenes (Newman) 3 X X
Neoperla carlsoni Stark & 1 X
Baumann
Neoperla clymene (Newman) 12 X X X X
Paragnetinafumosa (Banks) 1 X
Paragnetina kansensis (Banks) 1 X
Perlesta placida (Hagen) 20 X X X X X
Perlesta sp. A 1 X




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