ISSN 0870 - 3876 BOLETIM DO MUSEU MUNICIPAL DO FUNCHAL (HISTÓRIA NATURAL) Suplemento N.º 13

16th INTERNATIONAL CHIRONOMID SYMPOSIUM

SAMANTHA JANE HUGHES, MAHNAZ KADEM & MIGUEL ÂNGELO CARVALHO (Guest Editors)

Dezembro de 2008 FUNCHAL - MADEIRA

Editado pela Câmara Municipal do Funchal Composição: Museu Municipal do Funchal (História Natural) ,PSUHVVmRHDFDEDPHQWR2/LEHUDO(PSUHVDGH$UWHV*Ui¿FDV Foreword

This special supplement of the Boletim do Museu Municipal do Funchal contains extended abstracts, based upon communications given at the 16th International Chironomid Symposium which took place at the Casa da Luz Museum in Funchal, 25th - 28th July 2006, and was hosted and organized by the Centre for Macaronesian Studies (CEM) of the University of Madeira (UMa). The symposium provided an opportunity for chironomid researchers to attend sessions where communications of a remarkable scienti¿c level were given in Palaeolimnology, Biomonitoring, Toxicology & Biomonitoring, Ecology, , Morphology & Systematics, Physiology & Physiological Responses and Biogeography & Biodiversity. CEM included some ³¿UVWV´ in the organization of this international cycle of symposia such as a session on Palaeolimnology, emphasizing the importance of chironimids in assessing both past and present impacts in global issues such as climate change, an open debate on “Divulging Chironomid research: bibliography and data bases´ and a post symposium taxonomic workshop held at CEM’s laboratory facilities where researchers discussed and examined material and shared ideas. We extend our sincere thanks to everyone who contributed to the symposium’s success and to the preparation and publication of these proceedings, in particular the Director of the Museu Municipal do Funchal for agreeing to the publication of the symposium proceedings as a special supplement of the Boletim and colleagues from CEM and the Instituto Superior de Agronomia in Lisbon who translated proceedings abstracts into Portuguese. Many thanks to the organizing committee members for their dedication and willingness to carry out tasks prior to, during and after the symposium. Finally, a very special thank you to all of the sponsors for their extraordinary generosity, in particular the Empresa de Electricidade, various sectors of the Madeiran Regional Government and the Fundação para Ciência e a Tecnologia (FCT).

Samantha Jane Hughes Mahnaz Khadem President of the 16th International Chironomid Editor of the Proceedings of the 16th Symposium International Chironomid Symposium Chief Editor of the Proceedings of the 16th International Chironomid Symposium

Miguel Ângelo Carvalho President of CEM Sponsors of The 16th International Chironomid Symposium INDEX

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'DWHUHFHLYHG 2008 Proceedings of the 16th International Chironomid Symposium 21

COMPARISON AND SIGNIFICANCE OF CHIRONOMIDAE EMERGENCE FROM LAKE ERIE AND PRESQUE ISLE BAY, ERIE, PENNSYLVANIA, U. S. A.

By LEONARD C. FERRINGTON JR*1 EDWIN C. MASTELLER2 & JORGE A. SANTIAGO-BLAY3

With 1 Figure and 2 Tables

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Bol. Mus. Mun. Funchal, Sup. N.º 13 ,661 22 Boletim do Museu Municipal do Funchal (História Natural) Sup. No. 13

INTRODUCTION

Collections of surface-floating pupal exuviae (SFPE) are used in rapid bioassessment protocols (RBP) in both lotic (e. g., FERRINGTON 1987, FERRINGTON & CRISP 1989) and lentic habitats (e. g., RUSE 2002, RUFER & FERRINGTON 2008) to measure changes in community structure across water quality gradients. FERRINGTON et al.’s (1991) work on the HIÀFDFy, HIÀFLHQF\ and cost-effectiveness of SFPE to d-net collections found that the approach matched or exceeded d-net collections. Used on a monthly basis across a gradient of disturbed sites SFPE provided better resolution than a single set of d-net collections (SEALOCK & FERRINGTON 2008), similar to an RBP design recommended in BARBOUR et al. (1999). However, phenological emergence patterns meant that SFPE collections from June did not perform as well across the gradient as d-net collections. Consequently, it is necessary to better document the variability of results from SFPE collections at differing spatial and temporal scales that can be employed in RBP designs.

Lake Erie

Presque Isle Bay

Sample Sites

Figure 1. Location of sample sites on Lake Erie and Presque Isle Bay.

Presque Isle Bay (PIB) in Lake Erie is formed by Presque Isle, a large sand spit that extends approximately for 20 kilometres (Fig 1). PIB water and sediment quality has been contaminated by industrial and HIÁXHQW water discharges from municipal sewage treatment for over a century although pollution control and eIÁuent treatment have improved water quality in recent decades (DIZ 2005). However several contaminants associated with sediments of PIB may still be a concern. Although Lake Erie has been 2008 Proceedings of the 16th International Chironomid Symposium 

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MIDGE (DIPTERA: CHIRONOMIDAE AND CERATOPOGONIDAE) COMMUNITY RESPONSE TO CANAL DISCHARGE INTO EVERGLADES NATIONAL PARK, FLORIDA

By RICHARD E. JACOBSEN1

:LWK¿JXUHVDQGWDEOHV

ABSTRACT: Quantitative samples of chironomid and ceratopogonid pupal exuviae were collected along 4 nutrient gradients in Everglades National Park (ENP in order to determine midge community response to nutrient enrichment and identify possible indicators of water quality. Community abundance, species richness, and Shannon-Wiener diversity showed no consistent relationship with nutrient gradients. (LJKWVSHFLHVZHUHVLJQL¿FDQWO\VHQVLWLYHWRVRXUFHVRIHQULFKPHQWRIWKHVHVSHFLHV were also sensitive to nutrient enrichment in Water Conservation Area 2A (WCA-2A) VWXGLHGLQ6HYHQVSHFLHVZHUHVLJQL¿FDQWO\WROHUDQWWRDQGPRUHDEXQGDQWZLWK HQULFKPHQWEXWQRQHRIWKHVHVSHFLHVZHUHVLJQL¿FDQWO\WROHUDQWWRHQULFKPHQWLQ :&$$7KLVGLVFUHSDQF\LQWROHUDQWVSHFLHVSUREDEO\UHÀHFWVGLIIHUHQFHVLQVSHFLHV responses to low gradients in ENP versus the much steeper gradient in WCA-2A.

RESUMO: Com o objectivo de determinar a resposta das comunidades de DRHQULTXHFLPHQWRGHQXWULHQWHVHLGHQWL¿FDUHYHQWXDLVLQGLFDGRUHVGHTXDOLGDGHGD água, foram colhidas amostras quantitativas de exúvias de de quironomídeos HFHUDWRSRJRQtGHRVDRORQJRGHJUDGLHQWHVGHQXWULHQWHVSURYHQLHQWHVGHHÀXHQWHV de canais, no Parque Nacional de Everglades (PNE). A abundância da comunidade, ULTXH]DHVSHFt¿FDHRtQGLFHGHGLYHUVLGDGHGH6KDQQRQ:LHQHUQmRUHYHODUDPXPD UHODomRFRQVLVWHQWHFRPDSUR[LPLGDGHUHODWLYDDRVHÀXHQWHVGRVFDQDLV2LWRHVSpFLHV GHPRQVWUDUDPVHUVLJQL¿FDWLYDPHQWHPDLVVHQVtYHLVDRVHÀXHQWHVGRVFDQDLV'HVWDV IRUDPLJXDOPHQWHVHQVtYHLVDRHQULTXHFLPHQWRGHQXWULHQWHVQDÈUHDGH&RQVHUYDomRGD ÈJXD$ $&$$ HVWXGDGDHP6HWHHVSpFLHVUHYHODUDPVHUVLJQL¿FDWLYDPHQWH WROHUDQWHVHPDLVDEXQGDQWHVSHUWRGRVHÀXHQWHVGRVFDQDLVQRHQWDQWRQHQKXPD GHVWDVGHPRQVWURXVHUVLJQL¿FDWLYDPHQWHWROHUDQWHDRHQULTXHFLPHQWRQD$&$$ (VWDGLVFUHSkQFLDQDWROHUkQFLDUHÀHFWHSURYDYHOPHQWHGLIHUHQoDVQDUHVSRVWDGDV

1 IAP World Services and USGS-Florida Integrated Science Center, Everglades National Park, Homestead, Florida 33034 USA; E-mail: [email protected]

Bol. Mus. Mun. Funchal, Sup. N.º 13: 39-50, 2008 ISSN 0870-3876 40 Boletim do Museu Municipal do Funchal (História Natural) Sup. No. 13

HVSpFLHVDJUDGLHQWHVUHGX]LGRVQR31(HPFRQWUDSRVLomRDRVJUDGLHQWHVPXLWRPDLV acentuados observados na ACA-2.

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The Everglades is a phosphorus-limited oligotrophic marsh system that is highly sensitive to phosphorus enrichment (NOE et al. 'HVSLWHLWVQXWULHQWOLPLWDWLRQ the Everglades supports unusually high standing stocks of periphyton, a keystone feature of the system (TURNER et al., 1999). Marsh communities can undergo profound structural and functional changes to nutrient loads as low as 5 ppb above ambient levels (GAISER et al. 2005). However, the assimilation capacities of Everglades marshes for total phosphorus (TP) loading are so high that marsh waters near sources of enrichment often show no elevation in water TP. Agricultural and urban inputs of P currently threaten the biotic integrity of the Everglades ecosystem, particularly the northern Everglades where enriched canal water from agricultural areas causes eutrophication characterized by dense stands of Typha3URSRVHGPRGL¿FDWLRQVSODQQHGIRUWKH&HQWUDODQG6RXWK Florida Project as part of the Comprehensive Everglades Restoration Plan (CERP) will DOWHUZDWHUÀRZLQWR(YHUJODGHV1DWLRQDO3DUN (13 (IIHFWLYHELRPRQLWRULQJPHWKRGV QHHGWREHGHYHORSHGDQGLPSOHPHQWHGWRHQVXUHWKHVHPRGL¿FDWLRQVGRQRWGHJUDGH(13 marsh systems. Chironomid communities have been used for assessing water quality in the QRUWKHUQ(YHUJODGHV .,1*.,1* 5,&+$5'621 .,1*   OLVWHGFKLURQRPLGVSHFLHVWKDWZHUHLQGLFDWLYHRIVSHFL¿FQXWULHQWFRQGLWLRQVLQ:DWHU Conservation Area 2A (WCA-2A). However, his collections were conducted along a much greater nutrient gradient than those currently found within Everglades National Park (ENP). Many of the indicators that he found may be either absent or unresponsive to biological changes along nutrient gradients in ENP. Since ENP managers are interested LQLPSOHPHQWLQJPLGJHSXSDOH[XYLDHVDPSOLQJDVDHI¿FLHQWELRPRQLWRULQJPHWKRGIRU detecting nutrient enrichment, this study presents results of midge species and community UHVSRQVHVDORQJQXWULHQWJUDGLHQWVFUHDWHGE\FDQDOLQÀRZVLQWR(13DQGDWWHPSWV to identify community metrics and species that may serve as indicators of either water quality or nutrient enrichment in ENP marshes.

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0LGJHFRPPXQLWLHVZHUHVDPSOHGDORQJWUDQVHFWVIURPLQÀRZVLQWR(13/ 31W canal discharge into Taylor Slough, 332B retention pond discharge into the Rocky Glades, and from 2 Tamiami Canal culverts discharging into Northeast Shark River 6ORXJK 1(656  )LJ 7ZRKDELWDWW\SHVGH¿QHGE\WKHLUGRPLQDQWSODQWVSHFLHVZHUH sampled at 2 to 4 sample sites along each transect, Eleocharis and Cladium habitats were 2008 Proceedings of the 16th International Chironomid Symposium 41

sampled at 4 sites (50 m, 1.5 km, 4 km, and 8 km) downstream of discharge into Taylor Slough and at 3 sites along three transects in Shark Slough (100 m, 1.0 km, 3.0 km). At the S-332B retention pond, Cladium and Muhlenbergia habitats were sampled 50 m and PZHVWRIWKHSRQGRXWÀRZ4XDQWLWDWLYHVDPSOHVRIÀRDWLQJPLGJHSXSDOH[XYLDH were collected by skimming the water surface within four 0.25 m2 bottomless plastic corrals placed side-by-side within each habitat type, giving a 1.0 m2 total sample surface DUHD7KHGHQVLW\RIHPHUJHQWYHJHWDWLRQSHULSK\WRQDQGÀRDWLQJDOJDHLQ(YHUJODGHV marshes greatly inhibits exuvial drift; thus quantitative measures of exuvial species richness at the water surface were assumed to represent the density of emerging species per unit area of underlying marsh habitat. For each sample, we recorded water depth, the number of stems of each plant species present, and visual estimates of the percentage of ZDWHUVXUIDFHFRYHUHGE\ÀRDWLQJDOJDH PHWDSK\WRQ DQGVWHPVFRYHUHGE\FDOFDUHRXV HSLSK\WRQ&KLURQRPLGSXSDOH[XYLDHZHUHLGHQWL¿HGWRVSHFLHVRUPRUSKRVSHFLHVXVLQJ JACOBSEN (2008). Plant tissue, whole water, and soil samples were collected from each sampling site for nutrient analyses at the University of Florida Tropical Research and Educational Center, Homestead, FL. Total P of soil samples was determined by extraction using HCl.

Figure 1. Everglades National Park showing location of sampling transects downstream of canal discharges into the Park. 42 Boletim do Museu Municipal do Funchal (História Natural) Sup. No. 13

,QGLFDWRUVSHFLHVDQDO\VLV ,163$1'8)5Ç1( /(*(1'5( ZDV XVHGWRLGHQWLI\PLGJHWD[DZLWKVLJQL¿FDQWDI¿QLWLHVIRUPDUVKHVQHDURUIDUDZD\IURP FDQDORXWÀRZ,163$1ZDVFRQGXFWHGXVLQJ3&25' 0M06RIWZDUH*OHQHGHQ Beach, Oregon, USA). Separate INSPAN analyses were performed on samples from Eleocharis habitats, Cladium habits, and all habitats combined to increase the overall SRZHURIVLJQL¿FDQFHWHVWV

RESULTS

The nutrient gradients represented by these discharges were small (Fig. 2). Only water and EleocharisSODQWWLVVXHVDPSOHVVKRZHGLQFUHDVHVLQPHDQ3OHYHOVQHDULQÀRZ points (paired t-test: t=3.319, P=0.004; t=6.143, P=0.004 respectively). Water total-P OHYHOVZHUHKLJKHUQHDULQÀRZVDWERWKWUDQVHFWVVDPSOHGLQ1(656DQGQHDUUHWHQWLRQ pond 332B, but showed no change in Taylor slough near the L-31W canal. Eleocharis WLVVXHSHUFHQWWRWDO3FRQFHQWUDWLRQVZHUHKLJKHUQHDUFDQDORXÀRZLQWRXSSHU7D\ORU 6ORXJKDQG1(6561RVLJQL¿FDQWFKDQJHVZHUHREVHUYHGLQCladium tissue levels or VRLOOHYHOVRIWRWDO3ZLWKGLVWDQFHIURPLQÀRZV

Figure 2. Mean total-P (+SE) concentrations in water, soil, and plant tissue samples collected DORQJLQÀRZJUDGLHQWVVDPSOHGLQ(YHUJODGHV1DWLRQDO3DUN$XWXPQ$:DWHUVDPSOHV% Soil samples (HCl extraction). C. EleocharisWLVVXH'Cladium tissue. E. Comparison of water total-P gradients in ENP (this study) with those reported for WCA-2A by KING (2001). 2008 Proceedings of the 16th International Chironomid Symposium 43

Figure 3. Mean (+SE) abundance, taxa density, site species richness, and diversity of midge pupal H[XYLDHFROOHFWHGLQ$XWXPQVDPSOHVDORQJFDQDOLQÀRZJUDGLHQWVLQHDVWHUQ(YHUJODGHV National Park. A. Results from Cladium and Eleocharis habitats in northeast Shark River Slough DQG7D\ORU6ORXJKDORQJFDQDOLQÀRZJUDGLHQWV%5HVXOWVIURPCladium and Muhlenbergia KDELWDWVLQWKH5RFN\*ODGHVQHDULQÀRZVIURPUHWHQWLRQSRQG% 44 Boletim do Museu Municipal do Funchal (História Natural) Sup. No. 13

7RWDOH[XYLDHGHQVLW\GLGQRWFKDQJHZLWKSUR[LPLW\WRLQÀRZVLQHLWKHU7D\ORU Slough or NESRS, but increased near retention pond 332B (Fig. 3). Higher numbers of exuviae were collected in Eleocharis and Muhlenbergia habitats compared to Cladium habitats. Eleocharis habitats also had more periphyton growth (52±10% for Eleocharis versus 3±3% for Cladium, P (paired t-test) < 0.001) than Cladium habitats in sloughs. Species richness and diversity metrics showed no clear or consistent response UHODWLYHWRLQÀRZSUR[LPLW\Cladium samples tended to have higher mean species density than Eleocharis samples in sloughs and Muhlenbergia habitat near 332B. However, Cladium and Eleocharis habitat mean species densities did not consistently increase with proximity to canal discharges. Mean species density and site species richness increased near the L-31W canal in Taylor Slough and near the 332B retention pond. Cladium and Muhlenbergia habitats 50m west of retention pond 332B yielded a total of 44 taxa, including several species (e.g., Chironomus stigmaterus Say, Cladopelma forcipis (Rempel), Dicrotendipes modestus (Say)) considered to be indicators of enrichment $'$086 %5$1'7 +RZHYHUFRPPXQLWLHVLQ1(656VKRZHGQRLQFUHDVH LQVSHFLHVULFKQHVVQHDULQÀRZV6KDQQRQ:LHQHUGLYHUVLW\LQFUHDVHGVLJQL¿FDQWO\LQ CladiumVORXJKKDELWDWVQHDUFDQDOLQÀRZVEXWQRWLQEleocharis slough habitat and GURSSHGQHDU%RXWÀRZLQWKH5RFN\*ODGHV A total of 70 midge taxa were collected in samples from ENP (58 Chironomidae and 12 Ceratopogonidae; Table 1). Rocky Glades marshes near retention pond 332B had the largest number of species unique to its region (12 species, versus 3 species unique to Taylor Slough, and 7 species unique to NESRS), many of which are considered to be indicators of enrichment. (LJKWVSHFLHVZHUHVLJQL¿FDQWO\DVVRFLDWHGZLWKPDUVKHVIDUDZD\IURPFDQDO discharges and were considered to be sensitive to nutrient enrichment (Table 2). Seven of these species were also sensitive to phosphorus enrichment in WCA-2A (KING, 2001). Pseudochironomus articaudus Sæther is rare or absent in WCA-2A (JACOBSEN, 2008). Seven species, including the ceratopogonid, Bezzia cf. nobilisZHUHVLJQL¿FDQWO\ DVVRFLDWHGZLWKPDUVKHVQHDULQÀRZVDQGDUHFRQVLGHUHGWREHSRWHQWLDOLQGLFDWRUVRI enrichment for ENP. 2008 Proceedings of the 16th International Chironomid Symposium 45

TABLE 1. INSPAN analyses results for midge species collected along nutrient gradients in (YHUJODGHV1DWLRQDO3DUN1HDU QHDULQÀRZV WROHUDQWWRHQULFKPHQW )DU IDUIURPLQÀRZV (sensitive to enrichment); N = number of exuviae collected at near and far sites combined; IV = ,QGLFDWRUYDOXH SURGXFWRISUHYDOHQFHDQGSHUFHQWDEXQGDQFHDWSUHIHUUHGORFDWLRQ 6LJQL¿FDQWO\ associated species (P < 0.05) are indicated in bold lettering.

Preferred Relative location abundance in group (%) N IV P Chironomidae Tanypodinae Ablabesmyia sp. A Far 80 56 48.8 0.0029 Ablabesmyia sp. B Far 79 29 31.2 0.0478 Ablabesmyia sp. C Near 100 3 10.7 0.2374 Clinotanypus sp. Near 100 1 3.6 1 Fittkauimyia serta (Roback) Far 100 1 3.6 1 Labrundinia beckae Roback Near 100 1 3.6 1 Labrundinia maculata Roback Near 89 9 15.9 0.1353 Labrundinia neopilosella Beck & Beck Near 56 314 53.7 0.3548 Labrundinia sp. B Epler Near 100 1 3.6 1 Labrundinia sp. 6/10 Roback Near 100 2 3.6 1 Larsia decolorata (Malloch) Far 55 193 35.6 0.2481 Paramerina sp. Far 50 72 30.4 0.9179 Orthocladiinae Corynoneura sp. B Near 100 1 3.6 1 Limnophyes sp. Far 100 2 7.1 0.5021 Parakiefferiella coronata (Edwards) Far 67 189 47.6 0.0439 Pseudosmittia sp. Far 77 13 16.5 0.2108 Pseudochironomini Pseudochironomus articaudus Sæther Far 92 203 59.2 0.0003 Apedilum sp. Near 77 31 24.9 0.1123 Beardius breviculus Reiss & Sublette Far 54 267 29.1 0.2219 Beardius truncatus Reiss & Sublette Far 100 1 3.6 1 Chironomus stigmaterus Say Near 100 1 3.6 1 Chironomus sp. B Near 93 29 20.0 0.1180 Chironomus (Lobochironomus) sp. Near 97 31 20.7 0.0225 Cladopelma forcipis (Rempel) Near 100 3 10.7 0.2395 Cladopelma sp. A Near 58 31 18.7 0.4470 Cryptochironomus sp. B Near 73 15 13.1 0.4585 Dicrotendipes modestus (Say) Near 100 2 7.1 0.4930 Dicrotendipes simpsoni Epler Far 100 1 3.6 1 46 Boletim do Museu Municipal do Funchal (História Natural) Sup. No. 13

TABLE I (Cont.) Preferred Relative location abundance in group (%) N IV P Endochironomus nigricans (Johannsen) Far 67 3 4.8 1 Goeldichironomus cf.ÀXFWXDQVReiss Near 84 55 14.9 0.4631

Nilothauma sp. Far 96 69 58.1 0.0001 Parachironomus alatus (Beck) Far 100 2 7.1 0.4929 Parachironomus carinatus (Townes) Far 71 7 10.2 0.5034 Parachironomus sp. A Near 67 27 19 0.4806 Polypedilum beckae (Sublette) Near 93 98 29.8 0.0351 Polypedilum cf. falciforme Maschwitz Near 100 13 21.4 0.0241 Polypedilum simulans Townes Far 84 455 56.7 0.0161 Polypedilum trigonus Townes Near 52 118 34.5 0.8344 Polypedilum tritum (Walker) Near 93 41 29.8 0.0073 Polypedilum sp. K Near 50 2 1.8 1 Polypedilum sp. L Near 100 20 14.3 0.1083 Xenochironomus xenolabis (Kieffer) Far 100 5 10.7 0.2415 Zavreliella marmorata (Wulp) Near 72 163 28.2 0.6857 Tanytarsini Cladotanytarsus acornutus Jacobsen & Bilyj Far 95 65 40.9 0.0010 Cladotanytarsus sp. B Far 100 4 14.3 0.1141 Cladotanytarsus sp. C Far 82 656 70.3 0.0216 Tanytarsus limneticus Sublette Near 81 355 43.2 0.1524 Tanytarsus VS1' Near 100 17 32.1 0.0023 Paratanytarsus sp. B Near 100 1 3.6 1 Tanytarsus sp. A Near 76 38 16.4 0.4538 Tanytarsus sp. B Near 100 7 17.9 0.0488 Tanytarsus sp. C Near 100 5 14.3 0.1126 TanytarsusVS' VS5(SOHU  1HDU     Tanytarsus sp. E (= sp. J Epler) Near 100 2 7.1 0.4865 Tanytarsus sp. F Near 88 42 25.2 0.0540 Tanytarsus sp. G Near 56 524 49.8 0.4771 Tanytarsus sp. H Near 100 7 10.7 0.2326 Tanytarsus sp. I Near 50 6 3.6 1 Ceratopogonidae sp. A (=Dasyhelea cf. atlantis Wirth & Williams) Near 55 49 19.7 0.7314 sp. B (=Dasyhelea cf. major [Malloch]) Far 57 1852 57.2 0.3916 sp. C (=Bezzia cf. nobilis [Winnertz]) Near 90 208 80.7 0.0001 sp. E (=Stilobezzia sp.) Far 100 1 3.6 1 sp. G Far 76 42 29.9 0.0590 sp. H Near 100 1 3.6 1 sp. I Far 86 7 9.2 0.4869 sp. O Far 71 7 10.2 0.5170 sp. Q (=Alluaudomyia sp.) Far 100 3 10.7 0.2336 2008 Proceedings of the 16th International Chironomid Symposium 47

TABLE 2/LVWVRIPLGJHWD[DVLJQL¿FDQWO\DVVRFLDWHGZLWKORZQXWULHQWPDUVKHVDQGQXWULHQW enriched marshes in Everglades national Park (this study) and Water Conservation Area 2A (KING, 2001).

Indicators of low nutrient conditions ENP “nutrient-sensitive” WCA-2A “nutrient-sensitive” Ablabesmyia sp. A Ablabesmyia sp. A Ablabesmyia sp. B Ablabesmyia sp. B Parakiefferiella coronata Parakiefferiella coronata Nilothauma sp. Nilothauma sp. Polypedilum simulans Polypedilum simulans Cladotanytarsus acornutus Cladotanytarsus acornutus Cladotanytarsus sp. C Cladotanytarsus sp. C Pseudochironomus articaudus Parachironomus alatus Corynoneura sp. B Nanocladius alternantherae Beardius breviculus Paratanytarsus sp. B TanytarsusVS'

Indicators of nutrient enrichment ENP “nutrient-tolerant” WCA-2A “nutrient-tolerant” Chironomus (Loboch.) sp. Pseudochironomus richardsoni Polypedilum beckae Chironomus stigmaterus Polypedilum cf. falciforme Dicrotendipes modestus Polypedilum tritum Dicrotendipes simpsoni Tanytarsus sp. B Goeldichironomus holoprasinus TanytarsusVS1' Goeldichironomus cf. natans Bezzia cf. nobilis Kiefferulus sp. Polypedilum trigonum Tanytarsus sp. F Epler Tanytarsus sp. J Epler

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The unusual chemical and biotic features of the Florida Everglades (nutrient- limited, oligotrophic system, yet supporting the highest known standing stocks of periphyton in the world; TURNER et al., 1999) are quite distinct from other studied marsh systems. Increases in mean species density with nutrient enrichment were observed in VORXJKKDELWDWVE\5$'(5 5,&+$5'621  +RZHYHU.,1*  IRXQG 48 Boletim do Museu Municipal do Funchal (História Natural) Sup. No. 13

that species richness tended to show a unimodal, subsidy-stress response to enrichment in WCA-2A, and was unreliable as a measure of water quality. Further, he found that metrics of taxonomic structure and feeding ecology failed to show monotonic relationships with enrichment. McCORMICK et al. (2004) also observed dramatic shifts in species composition along the nutrient gradient in WCA-2A but no change in species diversity. 7KHVH¿QGLQJVVKRZWKDWVWUHDPELRDVVHVVPHQWPHWULFVDUHQRWHIIHFWLYHIRUDVVHVVLQJ Everglades marshes (TURNER et al., .,1*.,1* 5,&+$5'621  KLJKOLJKWLQJWKHQHHGWRXVHVSHFL¿FPHDVXUHVRIFRPPXQLW\VWUXFWXUHDQG indicator species groups for effective bioassessment of these distinct systems. Changes in community composition, particularly amongst species with different sensitivities to nutrient levels, is a more reliable method of assessing change in ENP water quality (KING, 2001). The large difference in the range of the WCA-2A and ENP nutrient gradients (Fig. 2E) probably accounts for the compositional differences between indicator species groups determined in this study versus KING (2001). Nutrient-sensitive species for ENP possibly represent a subset of WCA-2A nutrient–sensitive species that are particularly responsive to even small increases in nutrient loading (Table 2). Pseudochironomus articaudus has not been found in the northern Everglades (KING 2001; JACOBSEN, 2008). Nutrient-sensitive species in WCA-2A such as CorynoneuraVS'(3/(5   Nanocladius alternantherae 'HQG\ 6XEOHWWH Parachironomus alatus (Beck), and Paratanytarsus VS%(3/(5  ZHUHWRRUDUHLQWKLVVWXG\WRJHQHUDWHVLJQL¿FDQW indicator values. TanytarsusVS'VKRZHGRQO\DZHDNDI¿QLW\WRORZQXWULHQWPDUVKHV along the nutrient gradients in ENP. Likewise, ENP nutrient-tolerant species may be subsidized by the low levels of enrichment, but may become eliminated by conditions associated with the high P levels SUHVHQWLQ:&$$QHDUWKH+LOOVERURFDQDO1RQHRIWKHVSHFLHVVLJQL¿FDQWO\DVVRFLDWHG ZLWKPDUVKHVQHDU(13LQÀRZVDUHLQGLFDWRUVRIHQULFKPHQWLQ:&$$E\.,1*   Most species associated with enriched marshes in WCA-2A (Table 2) are well known LQGLFDWRUVRIHQULFKPHQW $'$086 %5$1'7 DQGDUHSUHVHQWLQ(13EXW overall water quality in the Park is high enough to ensure that they are rarely collected. Further collecting along nutrient gradients in ENP will be necessary to empirically validate these species as indicators of enriched waters. 6DPSOLQJSXSDHDQGH[XYLDHHQDEOHVVSHFLHVOHYHOLGHQWL¿FDWLRQRI&HUDWRSRJRQLGDH and their inclusion as water quality indicators. Ceratopogonids comprised from 25- 50% of all midge exuviae in samples (LISTON & TREXLER, 2005, and JACOBSEN, unpublished observations). In this study, Bezzia cf. nobilis was associated with ENP PDUVKHVQHDUFDQDOLQÀRZV2WKHUVLJQL¿FDQWLQGLFDWRUVRIZDWHUTXDOLW\PD\HYHQWXDOO\ be found in the species that mine within TyphaRU¿EURXVDOJDOPDWVLQHQULFKHGZDWHUVRU the abundant calcareous periphyton present in low-nutrient Everglades marshes (LISTON & TREXLER, 2005). 2008 Proceedings of the 16th International Chironomid Symposium 49

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%,%/,2*5$3+<

$'$0863 .%5$1'7 1990. Impacts on quality of inland wetlands of the United States: a survey of indicators, techniques, and applications of community level biomonitoring data. U.S. Environmental Protection Agency Report EPA/600/3-90/073, 406 pp.

'8)5Ç1(0 3/(*(1'5(   6SHFLHVDVVHPEODJHVDQGLQGLFDWRUVSHFLHVWKHQHHGIRUDÀH[LEOHDV\PPHWULFDODSSURDFK Ecological Monographs, 67: 345-366.

EPLER, J. H.: 2001. ,GHQWL¿FDWLRQ0DQXDOIRUWKH/DUYDO&KLURQRPLGDH 'LSWHUD RI1RUWKDQG6RXWK Carolina. A guide to the taxonomy of the of the southeastern United States, including Florida6SHFLDO3XEOLFDWLRQ6-631RUWK&DUROLQD'HSDUWPHQWRI(QYLURQPHQWDQG 1DWXUDO5HVRXUFHV5DOHLJK1&DQG6W-RKQV5LYHU:DWHU0DQDJHPHQW'LVWULFW3DODWND)/ 526 pp.

*$,6(5((-&75(;/(5-+5,&+$5'6'/&+,/'(56'/(($/(':$5'6/ -6&,172.-$<$&+$1'5$1*%12( 5'-21(6 2005. Cascading ecological effects of low level phosphorus enrichment in the Florida Everglades. Journal of Environmental Quality, 34: 717-723.

JACOBSEN, R. E.: 2008. A Key to the Pupal Exuviae of the Midges (Diptera:Chironomidae) of Everglades National Park, Florida.8QLWHG6WDWHV*HRORJLFDO6XUYH\6FLHQWL¿F,QYHVWLJDWLRQV5HSRUW 119 pp. 50 Boletim do Museu Municipal do Funchal (História Natural) Sup. No. 13

KING, R. S.: 2001. Dimensions of invertebrate assemblage organization in a phosphorus-limited Everglades landscape.3K'7KHVLV'XNH8QLYHUVLW\'XUKDP1&SS

.,1*56 &-5,&+$5'621 2002. Evaluating subsampling approaches and macroinvertebrate taxonomic resolution for wetland bioassessment. Journal of the North American Benthological Society, 21: 150-171.

LISTON, S. E., & J. C. TREXLER:   6SDWLDODQGWHPSRUDOVFDOLQJRIPDFURLQYHUWHEUDWHFRPPXQLWLHVLQKDELWLQJÀRDWLQJ periphyton mats in the Florida Everglades. Journal of the North American Benthological Society, 24: 832-844.

MCCORMICK396+8)25'5%(,,,DQG365$:/,. 2004. Changes in macroinvertebrate structure and function along a phosphorus gradient in the Florida Everglades. Hydrobiologia, 529: 113-132.

12(*%'/&+,/'(56$/(':$5'6((*$,6(5.-$<$&+$1'5$1'/((- 0(('(5-5,&+$5'6/-6&,172-&75(;/(5 5'-21(6 2001. Short-term changes in phosphorus storage in an oligotrophic Everglades wetland ecosystem receiving experimental nutrient enrichment. Biogeochemistry, 59: 239-267.

5$'(55% &-5,&+$5'621   5HVSRQVHRIPDFURLQYHUWHEUDWHVDQGVPDOO¿VKWRQXWULHQWHQULFKPHQWLQWKHQRUWKHUQ Everglades. Wetlands, 14: 134-146.

7851(5$075(;/(5-&-25'$1)6/$&.6-*(''(63 :/2)786 1999. Targeting ecosystem features for conservation: Standing crops in the Florida Everglades. Conservation Biology, 13: 898-911.

'DWHUHFHLYHG 2008 Proceedings of the 16th International Chironomid Symposium 51

BENTHIC MACROINVERTEBRATES AS WATER QUALITY INDICATORS IN ITALIAN LAKES

By L. MARZIALI1*, V. LENCIONI1, P. PARENTI2 & B. ROSSARO3

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A

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Fig. 1. A: Evenness (E), Shannon Wiener index (H), and taxa richness of the chironomid communities (average of 3 dates and standard errors). B: Indices of saprobity calculated from the chironomid and diatom communities, and indices of salinity (diatoms) at the stations studied (river km). 2008 Proceedings of the 16th International Chironomid Symposium 73

Fig. 2. Relative abundances of dominant species and conductivity values (all data averages of 3 sampling dates) at the river stations (river km); li = left shore, re = right shore.

DISCUSSION

The chironomid community of the river section studied was well characterized as “potamal” by typical taxonomic and ecological pUR¿les, but a clear ecological differentiation between sites could not be detected. This may UHÀHFW LQVLJQL¿FDQW differences in terms of both habitat structure (which would be of advantage to tracing chemical pollution in ¿HOG studies) and pollution qualities and quantities. Although the conductivity readings showed an apparently dramatic increase in salinity (Figs. 1, 2), the diatom salinity index indicates that this does not affect severely the functioning of the system. CASAS & VILCHEZ- QUERO (1996) found differences of chironomid communities in waters of different salinity, however, the conductivity values of the waters studied there were much higher than observed in this section of River Elbe. In this respect, the increasing abundances of N. dichromus and C. vanderwulpi may be due to other, unknown factors. These results suggest that the concentrations of pollutants stored in the upper layers of the sediment (HPA, 2005) apparently had no major LQÀXHQFH on community parameters despite remarkable point LQÀX[HV of serious pollutants. On the other hand, sediments sampled with corers smelled of petrol and other aromatics (esp. at site km 320), but obviuosly this had no effect on the chironomid community. This striking ¿QGLQJ may be explained by the contaminated sediments being covered with unpolluted sediments from upstream during ÀRRGV Probably, the pollutants are held in deeper sediment where they do not affect the benthic community living in the upper layers (down to 3 cm) or at the surface. Each spate may erode the sediment and cause a release of the stored pollutants. However, it is unclear whether this remobilisation of contaminated sediments affects the benthic community only locally or at other sites. Although chironomids reportedly are  %ROHWLPGR0XVHX0XQLFLSDOGR)XQFKDO +LVWyULD1DWXUDO  6XS1R

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'DWHUHFHLYHG ! 2008 Proceedings of the 16th International Chironomid Symposium 77

SAMPLING FREQUENCY REQUIRED FOR CHIRONOMID COMMUNITY RESOLUTION IN URBAN LAKES WITH CONTRASTING TROPHIC STATES.

By MORIYA MCGOVERN RUFER 1* & LEONARD C.FERRINGTON JR. 1

With 2 Tables and 3 Figures

$%675$&7:HIRXQGWKDWIRXUPLQXWHVDPSOHVRIFKLURQRPLGVXUIDFHÀRDWLQJ SXSDOH[XYLDHSHUVHDVRQZHUHVXI¿FLHQWWRFKDUDFWHUL]HRIWKHDEXQGDQW taxa in chironomid communities of urban lakes with contrasting trophic states (mean total epilimnetic phosphorus to mean lake depth [mld] of 1.1 – 133 mg/L/m). Generic richness decreased with increasing total mean phosphorus to mld.

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1 * corresponding author; [email protected] 'HSDUWPHQWRI(QWRPRORJ\8QLYHUVLW\RI0LQQHVRWD+RGVRQ+DOO)ROZHOO$YH6W3DXO01 86$7,  )D[  UXIHU#XPQHGXIHUUL#XPQ edu.

Bol. Mus. Mun. Funchal, Sup. N.º 13: 77-84, 2008 ISSN 0870-3876 78 %ROHWLPGR0XVHX0XQLFLSDOGR)XQFKDO +LVWyULD1DWXUDO  6XS1R

INTRODUCTION

7KHUHLVORQJKLVWRU\RIXVLQJFKLURQRPLGVWRDVVHVVODNHZDWHUTXDOLW\LQ(XURSH starting with 7+,(1(0$11LQDQGIROORZHGE\BRUNDIN  6$(7+(5  DQG:,('(5+2/0  DPRQJRWKHUV7KURXJKRXWWKHWKFHQWXU\DPRGHO of indicator species for lake trophic states has been developed (/,1'(*$$5' 6$(7+(5 ,QSDUWLFXODU6$(7+(5  DQG:,('(5+2/0  IRXQG chironomid communities to be related to the total mean epilimnetic phosphorus to mean lake depth ratio. This model has not been applied in the United States, particularly in urban areas of Minnesota. :HZDQWHGWRGHWHUPLQHDUHODWLYHO\IDVWLQH[SHQVLYHDQGHI¿FLHQWZD\WR PRQLWRUWKHUHVSRQVHRIELRWDWRZDWHUTXDOLW\LQODNHVRIWKH7ZLQ&LWLHV0HWURSROLWDQ Area (TCMA) of Minnesota, USA to be used by local lake managers. To determine ODNHWURSKLFVWDWXVXVLQJFKLURQRPLGVZH¿UVWQHHGHGWRGHWHUPLQHWKHVDPSOLQJHIIRUW UHTXLUHGWRUHVROYHWKHFKLURQRPLGFRPPXQLW\FRPSRVLWLRQ7KHUHIRUHWKHREMHFWLYH RIWKLVVWXG\ZDVWRGHWHUPLQHWKHVDPSOLQJIUHTXHQF\UHTXLUHGWRFROOHFWWKHDEXQGDQW taxa in urban lakes that can be used as indicators in a lake trophic model. We assessed FKLURQRPLGFRPPXQLWLHV7&0$ODNHVXVLQJVXUIDFHÀRDWLQJSXSDOH[XYLDH 6)3( VLQFH this method takes one-third the time to process than do chironomid larvae and allows for better taxonomic resolution (Ferrington et al. 

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,QWKH7&0$WKHUHDUHRYHUODNHV7KHVHODNHVZHUHIRUPHGDVWKHUHFHGLQJ JODFLHUVVFRXUHGWKHHDUWK¶VVXUIDFHDQGVXEVHTXHQWO\PHOWHGDIWHUWKHODVWLFHDJH7KH lakes are a valuable local resource for recreation, ecological habitat and the water supply system. The lakes in the TCMA span a gradient of mesotrophic to hypereutrophic, and we chose four mesotrophic lakes, eight eutrophic lakes and four hypereutrophic lakes for a total of 16 lakes (Table 1). We selected lakes (Fig. 1) based on the following criteria: 1) they do not have a known pollutant source other than organic enrichment, 2) they have ERDWDFFHVV RIWKHVXUURXQGLQJODQGXVHLVUHVLGHQWLDO WKH\DUHJHQHUDOO\ round in shape so that the lake has only one bay and consistent trophic state and pupal exuviae collect on the downwind end of the lake, 5) they are the same hydrologic type. :HFROOHFWHG&KLURQRPLGDH6)3( FERRINGTON et al. PRQWKO\GXULQJ the ice-free period of 2005 (April to November). Our method consisted of dipping a white SDQLQWRWKHZDWHURQWKHGRZQZLQGVKRUHRIWKHODNHWRFROOHFW6)3(7KHFRQWHQWVRI WKHSDQZHUHSRXUHGWKURXJKDPLFURQDSHUWXUHVLHYH7KLVWHFKQLTXHZDVUHSHDWHG IRUPLQXWHVDQGWKHVDPSOHZDVZDVKHGLQWRDMDUZLWKHWKDQRO¿HOGSUHVHUYDWLYH 2008 Proceedings of the 16th International Chironomid Symposium

Overall sampling consisted of 128 total samples (8 dates/lake x 16 lakes x 1 sample/lake/ date), collected over 2-3 consecutive days during the third week of each month. $IWHUHDFKPRQWKO\VDPSOHVHVVLRQWKH6)3(VDPSOHVZHUHSURFHVVHGWKHLQWKH ODE$FRXQWVXEVDPSOHRI6)3(ZDVUDQGRPO\SLFNHGXQGHU[PDJQL¿FDWLRQIURP D6\UDFXVHGLVKDQGLQWRDYLDORIHWKDQRO6DPSOHVFROOHFWHGLQ1RYHPEHUGLGQRW FRQWDLQDQ\6)3(DQGZHUHRPLWWHGIURPWKHDQDO\VLV6)3(ZHUHVOLGHPRXQWHGXQGHU [PDJQL¿FDWLRQXVLQJ(XSDUDOŠPRXQWLQJPHGLXPDQGLGHQWL¿HGWRJHQXVIROORZLQJ COFFMAN & FERRINGTON  DQG:,('(5+2/0   7KHGDWDZHUHDQDO\]HGE\GHWHUPLQLQJWKHQXPEHURIJHQHUDDQGWKHSURSRUWLRQ of total generic richness detected each month for each lake. The maximum detection HI¿FLHQF\ZDVWKHKLJKHVWSURSRUWLRQRIWRWDOJHQHULFULFKQHVVGHWHFWHGIRUHDFKODNHDQG what month it occurred. Each permutation of the proportion of total generic richness for 2 months, 3 months and 4 months was also calculated for each lake and again the PD[LPXPGHWHFWLRQHI¿FLHQF\ZDVWKHFRPELQDWLRQZLWKWKHKLJKHVWSURSRUWLRQRIWRWDO generic richness. Next, the generic richness for each lake was correlated to phosphorus to mean lake depth by linear regression.

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Elmo 1,12 15 13,4 41,8 83,4 4,3 43- M 35 &KULVWPDV       0  6TXDUH       0  /LWWOH/RQJ       0  Calhoun 2,5 27 10,6 27,4 170,4 3 54 - E 31 +DUULHW       (  6FKXW]       (  0F&DUURQ       (  Gervais 6,32 34 5,3 14,6 86,2 1,5 56 - E 27 Turtle 8 24 3 7,3 165,1 2,4 51 - E 33 2ZDVVR       (  &HQWHUYLOOH       (  &ROE\       1$+  /RULQJ       +  &HGDU       +  &RPR       +  80 Boletim do Museu Municipal do Funchal (História Natural) Sup. No. 13

Fig. 1. Study sites marked with black dots in the Minneapolis/St. Paul Metropolitan Area.

RESULTS

Over the course of the study 14,148 SFPE were collected with a mean of 875 exuviae per lake. The exuviae represented 51 total genera, with the range per lake from 20 to 35 genera. Ten genera (20%) were common to all 16 lakes. The chironomid community consisted of 9 Tanypodinae genera, 10 Orthocladiinae genera and 33 genera. Within Chironominae, there were 8 Tanytarsini genera, 1 Pseudochironomini genus and 24 Chironomini genera. For one sample date, maximum detection HI¿FLHQF\ ranged from 59% to 82% (Table 2). Two samples increased maximum detection HI¿FLHQF\ to 79% - 100%; three samples per season had a maximum detection HI¿FLHncy of 86% - 100%, and four samples per season ranged from 91% - 100% (Fig. 2). Six lakes (38%) had 100% detection HI¿FLHQF\ with four samples and an additional six lakes (38%) were only missing one taxon with four sample dates. The four remaining lakes were missing two taxa with four sample dates, yet two of these lakes had the highest generic richness and the other two yielded less specimens over the sample period than the mean exuviae collected per lake. The most genus-rich month per lake ranged from May to August. Eleven lakes (69%) had the highest richness in May or June, with the most lakes having the highest richness in June (Table 2). Generic richness decreased with increasing ratio of mean epilimnetic phosphorus/ mean lake depth, R2=0.56 (Fig. 3). Turtle lake (#10) had low total phosphorus (24mg/l), 2008 Proceedings of the 16th International Chironomid Symposium 81 but also a shallow mean depth (3m) which resulted in a phosphorus to mean depth ratio of 8mg/L/mld (Table1). Turtle lake was also the secondhighest in genericrichness (Table 1). When this lake was moved to ¿IWK in the order on Fig. 3 instead of tenth, the variance explained increased, R2=0.71.

TABLE 2. Proportion of total generic richness collected each month for each lake. The most genus-rich month is bolded; lakes are sorted by mean P/mean depth.

April May June July Aug. Sept. Oct. Elmo 0,15 0,590,59 0,38 0,56 0,47 0,21 Christmas 0,39 0,52 0,61 0,26 0,55 0,45 0,16 Square 0,16 0,28 0,81 0,75 0,56 0,38 0 Little Long 0,23 0,62 0,19 0,62 0,42 0,42 0 Calhoun 0,35 0,45 0,48 0,58 0,71 0,48 0 Harriet 0,26 0,52 0,74 0,44 0,59 0,59 0,22 Schutz 0,21 0,46 0,63 0,58 0,71 0,5 0,04 McCarron 0,11 0,39 0,57 0,64 0,32 0,43 0,11 Gervais 0,3 0,3 0,63 0,52 0,7 0,26 0 Turtle 0,09 0,61 0,67 0,64 0,45 0,45 0,06 Owasso 0,44 0,52 0,63 0,74 0,81 0,74 0 Centerville 0,5 0,57 0,64 0,82 0,25 0,29 0,07 Colby 0,25 0,67 0,75 0,5 0,71 0,21 0 Loring 0,09 0,74 0,17 0,43 0,52 0,39 0,13 Cedar 0,43 0,67 0,48 0,62 0,33 0,52 0 Como 0,48 0,65 0,35 0,48 0,57 0,22 0

100

90

80

70

60 1 sample

Percent of community detected Percent 2 samples 50 3 samples Fig. 2. Percent of the chironomid 4 samples communitc tydetectedwith1-4sample 40 dates during the ice-free season of 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 Lakes 2005 (April – October). 82 Boletim do Museu Municipal do Funchal (História Natural) Sup. No. 13

140 40

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25 80

R = 0.55805 20

60 Mean P/Mean Z 15 Generic Richness Mean P/Mean Depth 40 Generic Richness 10

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DISCUSSION

We believe we found arelativelysimple way for lake managers to determine the abundant taxa of the chironomid community present in urban lakes of contrasting trophic states. Collecting SFPE only takes 10 minutes effort per sample day per lake (FERRINGTON et al. 1991), and we found that four samples per lake per season are VXI¿FLHQW to characterize the abundant taxa of the chironomid community for the purpose of creating a model to determine trophic state. Four sample dates per season detected most of theabundant taxa in TCMA lakes (91%-100%). These urrban lakes ranged from hypereuttrophic (phosporus/mld 133 mg/l/m) to mesotrophic (phosporus/mld 1.1 mg/l/m). In all lakes, the April samples contained different taxa than May throughAugust sammples; therefore, werecommend sammpling four times a season, with one sample in April and three samples between May and September. If one only has the resources to sample three times a season, a majority of taxa will still be recovered (86%-100%). Forr a three month sample scheddule, we recommendd sampling April, June and July. For a two month sample schedule, we recommend sampling April and once between June and July. For just one sample a season, we recommend sampling in June. 2008 Proceedings of the 16th International Chironomid Symposium 83

As we expected, the generic richness was correlated with the phosphorus to mean lake depth ratio. Lakes with less phosphorus (mesotrophic and oligotrophic) are able to support a more heterogeneous chironomid community with a wider range of tolerances to organic enrichment. In lakes with more phosphorus (eutrophic and hypereutrophic), the intolerant taxa cannot be sustained and a more homogeneous, less diverse community is present. The aim of this study was to help lake managers and chironomidologists determine KRZPDQ\VDPSOHGDWHVDUHQHFHVVDU\WRFDWHJRUL]HDEXQGDQWWD[DRIWKHFRPPXQLW\XVLQJ 6)3(6LQFHDKLJKHUJHQHULFULFKQHVVLVSUHGLFWHGIRUROLJRWURSKLFODNHVRQHPD\QHHG more seasonal samples to detect a similar percentage of the total community.

ACKNOWLEDGEMENTS

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Date received: 06-04-2008 . 2008 Proceedings of the 16th International Chironomid Symposium 85

SAMPLING EFFICIENCY OF CHIRONOMIDAE (DIPTERA) ACROSS DISTURBANCE GRADIENTS

By ADAM W. S EALOCK*1 & LEONARD C. FERRINGTON JR2

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INTRODUCTION

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'DWHUHFHLYHG 2008 Proceedings of the 16th International Chironomid Symposium 141

DOES POLARIZED LIGHT GUIDE CHIRONOMIDS TO NAVIGATE TOWARD WATER SURFACES?

By NIKOLAY MELTSER1,YECHEZKEL KASHI2 AND MEIR BROZA 1*

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Styrofoam board ÀRDWLQJ near water HGJH In each observation day, traps were opened for 5 consecutive periods, 10 min. each. At the end of each 10 min. period PLGJHV were counted and removed. Statistical analysis. One-way ANOVA was used for Experiments: 1, 2 and 3A, while two-way ANOVA was used for experiment 4. For experiment 3B we used non- independent T-test.

RESULTS

Experiment I: Mosquitoes made a clear choice: KLJKHr mean numbers (76.9, 14.6 and 1.25) of rafts per barrel were collected from the three “black” treatments: Bc., Bex and B, respectively ¿J 1; P< 0.05 for all JURXSV  All black barrels differed VLJQL¿FDQWO\ from G (0.5).

A B

4 120 3.5

3 100 a 2.5 a 80 HJJPDVVHV a 2 a 60 of ʋRIHJJUDIWV 1.5 ʋ 40 1 Aver. b Aver. 0.5 c 20 c c 0 0 Treatments Treatments B.c. B.ex. B. G. B.c B.ex B. G.

)LJ 1. Mean number of chironomid HJJPDVVHV (A) and culicid rafts (B), Elroi 6SULQJ Bars with same letter are not VLJQL¿FDQWO\ different. A P<0.001. B P< 0.05. (B) Black barrel/ tap water. (Bc) black cane plant added. Black (Bex) extract of cane added. Green (G) JUHHQ barrels with tap water.

The low number of Chironomid eJJ masses indicated that total chironomid population at this place was quite low but results indicate that females responded equally to all three black choices (B.c. 1.58; B.ex. 1.54; B. 1.96 (P>0.05)), but avoided the JUHHQ barrels (G; 0.2). For all black and JUHHQ barrels (P< 0.05). Female mosquitoes responded both to the chemical nature of water and the barrel colour. 1RQELWLQJ PLGJHV responded to the visual stimulus only in present conditions. Experiment II. The next experiments were carried out in Tiv’on Waste Stabilization Ponds (WSP), a habitat with hiJK population levels of Chironomidae. Results are summarized in )LJ 2. In this ¿JXUH are shown observations in a typical midseason month. Oviposition was VLJQL¿FDQWO\ KLJKHU (P<005) in the black (21.5) than in the JUHHQ (7.44) barrels, and in the JUHHQ than in the white (1.0) barrels. Early in the season, despite the limited number of HJJ masses, the attraction of the different barrels 2008 Proceedings of the 16th International Chironomid Symposium 145 remained as above. In fact, the least attractive barrel, W (white), almost did not attract any females (Fig. 2A).

35

30 a 25

20

of egg-masses 15

ʋ b 10

Aver. 5 c

0 Treatments

Black Green White

Fig. 2. Experiment near Waste Stabilization Ponds (WSP). Mean Number of egg-mass in - Black, Green and White barrels with tap water. Bars with different letters are VLJQL¿FDQWO\ different (P< 0.001).

Experiment III. Barrels were ¿lled with diluted (1:3) oxidation pond water. Euglena sp., the green unicellular alga, was the dominant phytoplankton (almost exclusive). Algae density on day 2 post-dilution was 7.5*103 cells/L., but by day 11 it had reached these levels: B, black: 1.5 * 104 / ml; G, green: 1.5 * 105 / ml; S, silver: 2.5 * 105 / ml.

A B

18 14

16 12 14

a 10 a 12

10 8 a a 8 of egg-masses

of egg-masses 6 ʋ

6 ʋ b 4 4 Aver. Aver.

2 c 2

0 Treatments 0 Treatments Black Green Silvered Black Green Silvered

Fig. 3. Oxidation pond water experiment. Mean Number of egg-masses/day collected in Black, Green and silvered barrels ¿OOHG with diluted (1:3) pond water. A data on day 2 (diluted phytoplankton) P <0.05). B day 11, during phytoplankton blooming, P> 0.05. Bars with same letter DUHQRWVLJQL¿FDQWO\GLfferent. 146 Boletim do Museu Municipal do Funchal (História Natural) Sup. No. 13

The load of egg-masses collected on days 2 and 11 of the experiment are shown in Fig. 3A and 3B. In 3A it is seen that females made a distinct choice in favor of the black barrels (21.6) rather than the green (6.9), and only 1.6 of the silver (P< 0.05). After 9 days all barrels yielded similar numbers of egg masses (P> 0.05). Moreover, shiny silver barrels attracted even higher numbers of ovipositing females (18.25) than the other two: black (13.5) and green (12.25). At this stage light rays were UHÀHFWHG from the shiny bottom and wall above and below the water surface (on S) and were non-polarized. Intensity of this light was higher than intensity of polarized light reÀected from the water surface. It reduced the effect of polarization. The numbers of laid eggs descended in the order black > green > white. Ten days later (Fig. 3B) phytoplankton reached the blooming stage, which was more intensive in the shiny silver barrels that absorbed maximum sun radiation (Fig. 4). Results indicate that at the early stage (Fig. 3A) females responded to light UHÀHFWHG from the barrel walls according to its colour, at the second stage (Fig. 3B) females’YLVLRQUHVSRQGHGWRWKHSRODUL]HGUHÀHFWHGZDWHUVXUIDFHUD\V

A

B 1

4 1

3

2

Fig. 4. A. Clean water with very low concentration of algae. All rays were UHÀHFWHG from the shiny walls and bottom of barrel are non polarized. B. Silvered barrel, ¿OOHG with pond water after development of Euglena algae en masse. (1) Non-polarized rays hits water surface and barrel walls above and below water. (2) Dark shiny water surface UHÀHFWHG horizontally polarized light. (3) Rays inserted into barrel depth absorbed by dark water. (4) Rays hit the aluminum foil above ZDWHUVXUIDFHUHÀHFWHGZLWKRXWSRODUL]DWLRQ

Experiment IV. In the prior experiments we con¿UPHd that differential UHÀHFWLon of light through the different treatments/colours support females’ site selection. However, although we assumed that this was related to sensing horizontal polarized light we could not exclude light intensity as being important. This question was resolved at the next stage (Fig 4). In the experiment with light traps VLJQL¿FDQWO\ higher numbers of females were captured in polarized than non-polarized traps ¿g 4; mean numbers: 42.0 and 25.75). The 2008 Proceedings of the 16th International Chironomid Symposium 147

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'DWHUHFHLYHG CHIRONOMIDAE (DIPTERA) IN ALPINE LAKES: A STUDY OF SUBFOSSIL ASSEMBLAGES IN LAKE SURFACE SEDIMENTS

By OLIVER HEIRI1* & ANDRÉ F. LOTTER1

With 3 Figures and 1 Table

ABSTRACT: Subfossil chironomid assemblages in surface sediments from Swiss PRXQWDLQODNHVVKRZDVWURQJVWDWLVWLFDOO\VLJQL¿FDQWUHODWLRQVKLSZLWKODNHGHSWK DQGZLWKSDUDPHWHUVUHODWHGWRDOWLWXGHVXFKDV-XO\WHPSHUDWXUHGLVVROYHGRUJDQLF FDUERQFRQFHQWUDWLRQRURUJDQLFPDWWHUFRQWHQWRIWKHVHGLPHQWV8QH[SHFWHGO\ZDWHU FKHPLVWU\YDULDEOHVVXFKDVWRWDOSKRVSKRUXVFRQFHQWUDWLRQDQGS+ZHUHQRWLGHQWL¿HG DVUHOHYDQWSDUDPHWHUV

RESUMO: As comunidades dos sub-fósseis quironomídeos existentes nos VHGLPHQWRVGHVXSHUItFLHSUHVHQWHVQRVODJRVPRQWDQKRVRVGD6XLoDPRVWUDUDPXPD DVVRFLDomRHVWDWLVWLFDPHQWHVLJQL¿FDWLYDFRPXPDVpULHGHSDUkPHWURVDPELHQWDLV 6mRGHGHVWDFDUDSURIXQGLGDGHGRODJRPDVWDPEpPSDUkPHWURVUHODFLRQDGRVFRPD DOWLWXGHWDOFRPRDWHPSHUDWXUDGRPrVGH-XOKRFRQFHQWUDomRGHFDUERQRRUJkQLFR GLVVROYLGRHFRQWH~GRGHPDWpULDRUJkQLFDGRVVHGLPHQWRV7RGDYLDHDSHVDUGH HVSHUDGRDVYDULiYHLVUHODFLRQDGDVFRPDTXtPLFDGDiJXDWDOFRPRFRQFHQWUDomRGH IyVIRURHS+QmRIRUDPFRQVLGHUDGDVFRPRYDULiYHLVHVWDWLVWLFDPHQWHUHOHYDQWHV

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for fossil chironomids using standard methods. Further details on ¿eldwork and the measurement of environmental variables are presented in LOTTER et al. (1997), HEIRI (2001) and BIGLER et al. (2006). Of the 115 chironomid assemblages presented in these publications only those from 45 mountain lakes situated higher than 1800 m a.s.l. are discussed here (Fig. 1). For direct gradient analyses we used the program CANOCO version 4.5 (TER BRAAK & ŠMILAUER 1998).

Figure 1. Outline map of Switzerland. The circles indicate the different study sites.

RESULTS AND DISCUSSION

The distribution of most chironomid taxa showed a strong relationship with altitude (Fig. 2). Lakes higher than 2400 m a.s.l. were dominated by taxa such as Pseudodiamesa, Micropsectra radialis-type and Paracladius. Between ca. 2400 and 2000 m a.s.l. taxa such as M. radialis-type, Psectrocladius sordidellus-type, and Tanytarsus lugens-type showed high abundances and chironomid remains belonging to, e.g., Paratanytarsus austriacus-type, Heterotrissocladius marcidus-type, Zavrelimyia type A and Micropsectra insignilobus/contracta-type reached moderately high abundances. In lakes between 1800 and 2000 m a.s.l. taxa such as T. lugens-type, H. marcidus-type, Chironomus anthracinus- type and Stictochironomus were found at high abundances. Furthermore, a range of taxa which are absent or only occur at low abundances at higher altitudes occur regularly in these lakes, e.g. Dicrotendipes nervosus-type, Heterotrissocladius grimshawi/scutellatus- type, or Microtendipes. 180 Boletim do Museu Municipal do Funchal (História Natural) Sup. No. 13

We used direct gradient analysis (Table 1) to assess how well the measured environmental variables can explain the distribution of chironomid remains in the study lakes. Variables strongly correlated with altitude all explained statistically VLJQL¿FDQW amounts of variance in the chironomid assemblages. Similar to ¿ndings by BIGLER et al. (2006), dissolved organic carbon of the lake water (DOC) and the organic content of the sediments explained more variance than July air temperature. Dissolved or total organic carbon concentration of lake water and the organic matter content of the sediments were also identi¿ed as important explanatory variables for chironomid assemblages in subarctic and arctic regions. It therefore seems that at the cold end of the temperature gradient the relationship between chironomid assemblages and summer temperature that has been reported by numerous studies may be iQÀuenced by the effects of organic carbon content of lake water and sediments on the larvae. In addition to parameters correlated with altitude, maximum water depth was also LGHQWL¿HG as a VLJQL¿FDQW explanatory variable (Table 1). Again, this is not surprising, since the depth of a lake basin will have a distinct LQÀXHQFH on factors of importance for the survival of chironomid larvae, e.g., water temperature at the sediment-water interface, light and food availability, and habitat structure (e.g. composition of the substrate, presence or absence of aquatic macrophytes).

Fig. 2: Subfossil chironomid assemblages analyzed in the surface sediments of 45 mountain lakes in Switzerland. The sites are arranged according to altitude and only the most abundant taxa are shown. 2008 Proceedings of the 16th International Chironomid Symposium 181

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Environmental parameter Range n r % Var $OWLWXGH PDVO      0HDQ-XO\DLUWHPSHUDWXUH ƒ&      0D[ZDWHUGHSWK P      'LVVROYHGRUJDQLFFDUERQ '2& FRQFHQWUDWLRQ PJO      $ONDOLQLW\ PPROO      S+     7RWDOSKRVSKRUXV 73 FRQFHQWUDWLRQ —JO      6HGLPHQWDU\RUJDQLFPDWWHU ':     

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Fig. 3. Distribution of selected chironomid taxa in the 45 study lakes with respect to summer temperature and DOC concentrations (A-B), maximum water depth (C-E), and the organic content of the sediment (F). The symbol size in the different scatter plots indicates the percent abundance of the chironomid taxa in the lakes. 2008 Proceedings of the 16th International Chironomid Symposium 183

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CHIRONOMID-BASED INFERENCES OF LOCAL AND REGIONAL ENVIRONMENTAL CHANGE DURING THE EARLY MIDDLE WEICHSELIAN IN NORTHEAST FINLAND

By STEFAN ENGELS1*, SJOERD J.P. BOHNCKE1,JOHANNA A.A. BOS1, KARIN HELMENS2 & OLIVER HEIRI3

With 2 Figures

ABSTRACT: A fossil-rich lacustrine deposit dated to ca. 50 thousand years ago with little post-depositional disturbance was recovered from the high-latitude Sokli- site (NE Finland). Analysis of fossil chironomid remains showed that a shallow lake was present at the study site, and that climate conditions were dynamic during the lake’s existence.

RESUMO: Um depósito lacustre rico em fósseis, aproximadamente com 50 mil anos de idade, foi recuperado numa área a grande latitude no Norte da Finlândia (Sokli). O presente depósito apresentava pouca perturbação pós-deposicão. As análises de restos dos fósseis quironomídeos revelou que o depósito se encontrava num lago de pouco profundidade, tendo estado sujeito a condições climáticas dinâmicas.

1.Department of Palaeoclimatology and Geomorphology, Vrije Universiteit Amsterdam, De Boelelaan 1085, 1081 HV Amsterdam, The Netherlands, [email protected]; [email protected]; [email protected] (*Author for correspondence) 2. Department of Physical Geography & Quaternary Geology, Stockholm University, SE-106 91 Stockholm, Sweden, [email protected] 3. Palaeoecology, Institute of Environmental Biology, Faculty of Science, Utrecht University, Laboratory of Palaeobotany and Palynology, Budapestlaan 4, 3584 CD Utrecht, The Netherlands, o.m.heiri@bio. uu.nl

Bol. Mus. Mun. Funchal, Sup. N.º 13: 185-190, 2008 ISSN 0870-3876 186 Boletim do Museu Municipal do Funchal (História Natural) Sup. No. 13

INTRODUCTION

Fossil chironomids have been widely used to reconstruct lacustrine environments and regional climate conditions during the Late-Glacial and Holocene periods. However, chironomid-based palaeoenvironmental reconstructions covering periods prior to ca. 15,000 years before present (BP) are still sparse (BROOKS, 2006). An exceptional lacustrine record dating back to the earlier part of the Middle Weichselian, ca 50 thousand years BP, was obtained from the Sokli site in Northeast Finland (HELMENS et al., 2000; 2007; Fig 1). Different sedimentological and botanical proxy-indicators were analyzed in order to reconstruct local and regional environmental changes. Furthermore, fossil chironomid remains were studied in the sequence to examine palaeoenvironmental changes and to reconstruct the LQÀXHQFH of external factors such as climate change on the aquatic ecosystem.

The Sokli site A 26m long and nearly continuous sedimentary sequence spanning the last 130,000 years has been encountered in a small sedimentary basin (67º48’N, 29º18’E, 220m a.s.l.) in Northeast Finland (see Fig. 1). Several glacial-deglacial cycles can be recognized in the sedimentary record, and 4 successive lacustrine bodies have been identi¿ed (HELMENS et al., 2000; 2007). Using radiocarbon and luminescence dating, the lower 3 lacustrine deposits were dated to the last interglacial (equivalent to MIS-5E, ca 120 ka BP), the Brørup interstadial (equivalent to MIS- 5C, ca 100 ka BP) and the Odderade interstadial (equivalent to MIS-5A, ca 80 ka BP). The ¿UVt results of a high-resolution fossil analysis on the uppermost limnic deposit are presented here. These sediments have been dated to the earlier part of the middle Weichselian, ca. 50 ka BP.

Fig. 1. Map of Finland, showing the location of the Sokli study site. 2008 Proceedings of the 16th International Chironomid Symposium 187

METHODS

The interval between 5.20 and 7.20 m core depth (in borehole Sokli B-series) was subsampled in 5-centimeter thick intervals and 39 wet sediment samples (weight range: 3.65 – 38.54 g) were used for chironomid analysis. First, the samples were immersed in 10% KOH for at least 4 hours and subsequently rinsed on a 100 µm sieve to remove ¿QH particles from the samples. Using a dissecting microscope, chironomid remains were hand picked at 35x PDJQL¿FDWLRQ and mounted on permanent slides. The fossil head capsules ZHUHLGHQWL¿HGIROORZLQJWKHOLWHUDWXUHOLVWHGLQ%2+1&.(et al. (2008).

Chironomid record and environmental inferences A total of 65 chironomid taxa were identi¿ed in the sediments of the Sokli-record. In the lowermost part of the record chironomid remains were sparse, but above 6.75 m core depth, a minimum count sum of 50 head capsules (hc) was reached for all samples. The average count sum in the samples above 6.75m core depth was 89 hc/sample (range: 54 – 199 hc). The high number of taxa with a preference for littoral to sub-littoral habitats suggests that throughout the record a shallow lake was present at the Sokli site. The lowermost samples (up to 6.75m core depth; chironomid-zone I) represent the pioneer stage of the record. Different species are present, but as counts are low in this part of the record, the abundances of the encountered taxa are highly variable (Figure 2). Among the taxa that ¿UVW appeared at the Sokli-site were Tanytarsus lugens- type, Microtendipes and Polypedilum nubeculosum-type.

Fig 2: &hironomid diagram for the Sokli study site showing selected taxa and zone boundaries. Taxa are expressed as a percentage of the total head capsule number, the chironomid productivity as the number of head capsules per gram of wet sediment. 188 Boletim do Museu Municipal do Funchal (História Natural) Sup. No. 13

The onset of chironomid-zone II shows a sharp increase in the concentration of chironomid remains, reaching values of 50 hc/g wet sediment. The assemblages are initially dominated by Chironomus anthracinus-type and Procladius. Both these taxa have a large temperature tolerance, but i.e. HEIRI (2001) states that C. anthracinus-type reaches its highest modern abundances in lakes at intermediate elevation in the Swiss Alps, rather than in lowland lakes. Towards the upper part of this zone, the abundance of T. lugens-type increases. This taxon also has a large temperature-tolerance, but has been considered to be indicative for cool and oligotrophic conditions. Toward the top of chironomid-zone II, abundances of C. anthracinus-type, Procladius and T. lugens- type decline, and these taxa are replaced by taxa that are indicative of higher July air temperatures such as Polypedilum nubeculosum-type and Cladotanytarsus type I. This gradual transition could be the result of a steadily increasing summer temperature at the Sokli-site. The peak of Endochironomus albipennis-type near the end of chironomid zone II might indicate rising temperatures too, but also could be related to increased nutrient availability in the lake. Cladotanytarsus type I is the dominant taxon at the onset of chironomid-zone III, showing a decreasing abundance over the entire zone. Another important taxon in this zone is Microtendipes, which reaches a maximum abundance of 20%. BEDFORD et al. (2004) state that this genus is “generally considered to be a warm stenothermic taxon … but at Whitrig Bog this taxon responded best to intermediate temperatures”. The increase in Microtendipes FRXOGWKHUHIRUHEHLQGLFDWLYHRIWHPSHUDWXUHVWKDWDUHEHVWFODVVL¿HGDV “intermediate to warm”, and the gradual warming trend that was observed in chironomid zone II is probably interrupted. The absence of taxa indicative of high nutrient availability, together with the relatively low concentration of chironomid remains, suggests a return to mesotrophic conditions. 'XULQJWKH¿QDOVWDJHRIWKHODNH FKLURQRPLG]RQH,9 Microtendipes disappears from the spectrum and is replaced by Corynocera ambigua. This species was formerly considered to be a true cold-water stenotherm, but those ideas have recently been questioned by BRODERSEN & LINDEGAARD (1999). Although the adult midges of C. ambigua DUHKDUGO\DEOHWRÀ\WKH\KDYHEHHQVKRZQWREHIDVWPLJUDWLQJDQG the temperature-preference of this species is probably “intermediate” to “low-arctic”. Paracladius, a taxon that is generally considered to be a true cold-stenotherm, shows two phases with high abundances: around 5.90m core depth and in the uppermost part of the core, between 5.20-5.50m. Thirdly, Stictochironomus shows high values in the upper part of the record. All these changes together can be interpreted as indicating a return to colder conditions. However, Polypedilum nubeculosum-type and Cladotanytarsus type- I are still the dominant taxa in the assemblages, and they imply intermediate- to warm summer temperatures. An interesting feature of chironomid zone IV is the occurrence of stream- indicating taxa such as Eukiefferiella and Prodiamesa. Together with the coarser-grained 2008 Proceedings of the 16th International Chironomid Symposium 189

VHGLPHQWLQWKLVLQWHUYDOWKLVVXJJHVWVLQFUHDVLQJLQÀXHQFHRIWKHQHDUE\LFHVKHHWRQWKH lake-catchment through proglacial streams. The close and advancing ice sheet is further demonstrated by the uppermost part of the lacustrine deposit, which gradually changes into a glacial till deposit, without a hiatus in the sedimentary sequence (HELMENS et al., 2007).

CONCLUSIONS

A fossil-rich lacustrine deposit dated back to the ca. 50ka ago with little post- depositional disturbance was recovered from the high-latitude Sokli-site (NE Finland). Chironomid analysis of this sequence showed changing concentrations of chironomid remains and a dynamic composition of the chironomid- assemblages throughout the record: ‡ 7KHKLJKQXPEHURIWD[DZLWKDSUHIHUHQFHIRUOLWWRUDOWRVXEOLWWRUDOKDELWDWVVXJJHVWV that a shallow lake must have existed at the Sokli site. ‡ 7KHFKLURQRPLGDVVHPEODJHVDQGWKHFKDQJHVEHWZHHQWKHGLIIHUHQW]RQHVVXJJHVW that climate was a major forcing factor with respect to the presence of the different chironomid taxa. The Sokli-site therefore offers the possibility to derive quantitative palaeotemperature estimates based on fossil chironomid remains, for a time period where the number of such records is still very limited.

REFERENCES

BOHNCKE, S.J.P., J.A.A. BOS, S. ENGELS, O. HEIRI & C. KASSE: 2008. Rapid climatic events as recorded in Middle Weichselian thermokarst lake sediments. Quaternary Science Reviews 27: 162-174.

BEDFORD, A., R.J. JONES, B. LANG, S.J. BROOKS & J.D. MARSHALL: 2004. A Late-Glacial chironomid record from Hawes Water, Northwest England. Journal of Quaternary Science 19: 281-290.

BRODERSEN, K.P. & C. LINDEGAARD: 1999. Mass occurrence and sporadic distribution of Corynocera ambigua Zetterstedt (Diptera, Chironomidae) in Danish Lakes. Neo- and palaeolimnological records. Journal of Paleolimnology 22: 41-52.

BROOKS, S.J: 2006. Fossil midges (Diptera: Chironomidae) as palaeoclimatic indicators for the Eurasian region. Quaternary Science Reviews 25: 1894-1910. 190 Boletim do Museu Municipal do Funchal (História Natural) Sup. No. 13

HEIRI, O: 2001. Holocene palaeolimnology of Swiss mountain lakes reconstructed using subfossil chironomid remains: past climate and prehistoric human impact on lake ecosystems. Ph.D. Thesis, University of Bern, Switzerland.

HELMENS, K.F., M.E. RÄSÄNEN, P.W. JOHANSSON, H. JUNGNER & K. KORJONEN: 2000. The Last Interglacial-Glacial cycle in NE Fennoscandia: a nearly continuous record from Sokli (Finnish Lapland). Quaternary Science Reviews 19: 1605-1623.

HELMENS, K.F., P.W. JOHANSSON, M.E. RÄSÄNEN, H. ALEXANDERSON & K.O. ESKOLA: 2007. A series of ice-free intervals continuing into MIS 3 recorded in the central area of Fennoscandian glaciation: late Quaternary climate-stratigraphy at Sokli. Geological Society of Finland Bulletin 79: 17-39.

Date received: 06-04-2008. 2008 Proceedings of the 16th International Chironomid Symposium 191

DETERMINATION OF CHIRONOMIDAE THERMAL PREFERENCES AND THERMAL PARTITIONING AMONG CLOSELY RELATED TAXA IN A MINNESOTA STREAM USING SURFACE FLOATING PUPAL EXUVIAE

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INTRODUCTION

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RESULTS

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2003. Chironominae was the most diverse subfamily with 60 species followed by Orthocladiinae with 48 species. Less diverse were the Tanypodinae with 19 species and the Diamesinae with one species identi¿ed. Mean emergence temperatures for subfamilies ranged from 5.32 to 20.19°C with Diamesinae emerging at the lowest temperatures and Tanypodinae at the highest (Fig. 1). Orthocladiinae and Chironominae had mean emergence temperatures of 16.83 and 18.48°C, respectively. Mean emergence temperatures for genera ranged from 5.32°C (Diamesa) to 24.52°C (Paramerina) (Fig. 2). At lower temperatures there was a greater prevalence of Orthocladiinae genera, but at warmer temperatures, Chironominae and Tanypodinae genera were more common (Figure 2). There were a few notable exceptions to these generalizations with Micropsectra emerging on average at 11.71°C and Corynoneura emerging on average at 22.51°C. Several other genera also did not match subfamily predictions (e.g. Glyptotendipes, Parachironomus, Epoicocladius, Eukiefferiella; Fig. 2), but the small number of specimens collected for these genera cause some uncertainty regarding their thermal preferences.

Fig. 1. Violin plots of subfamily relative abundance at emergence temperatures (plot width = relative abundance; line = median, and solid circle is mean).

The thermal preference ranges for both natural and simulated genera increased from smaller to larger genera (Fig. 3). For two, three, and four species genera, simulated genera had greater thermal preference ranges. In contrast, natural genera had similar thermal preference ranges for ¿Ye species genera and greater ranges for ten species genera when compared to simulated genera. The thermal preference ranges for natural two and three species genera were more variable than other genera sizes and treatments. This larger variation was the result of one genus among the two species genera and three genera among three species genera which had considerably larger thermal preferences ranges than the other genera with similar genus sizes. 2008 Proceedings of the 16th International Chironomid Symposium 195

Fig. 2. Violin plots of relative abundance for genera at emergence temperatures (plot width = relative abundance; line = median; solid circle = mean; dark grey = Diamesinae; medium grey = Orthocladiinae; light grey = Chironominae; white = Tanypodinae).

Fig. 3. Box plots of thermal preference ranges for natural and simulated genera of different sizes (middle line = median; upper and lower bounds of the box = 25th and 75th percentiles; whisker caps = 10th and 90th percentiles; circles = outliers). 196 %ROHWLPGR0XVHX0XQLFLSDOGR)XQFKDO +LVWyULD1DWXUDO  6XS1R

DISCUSSION

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ACKNOWLEDGMENTS

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BIBLIOGRAPHY

BECKETT, D.C.:  3KHQRORJ\RIODUYDO&KLURQRPLGDHRIDODUJHWHPSHUDWH1HDUFWLFULYHUJournal of Freshwater Ecology 7: 303-316. 198 %ROHWLPGR0XVHX0XQLFLSDOGR)XQFKDO +LVWyULD1DWXUDO  6XS1R

COFFMAN, W.P. :  )DFWRUVWKDQGHWHUPLQHWKHVSHFLHVULFKQHVVRIORWLFFRPPXQLWLHVRI&KLURQRPLGDHActa Biologica Debrecen Oecologica Hungarica 3

COFFMAN, W.P. & C.L. DE LA ROSA:  7D[RQRPLFFRPSRVLWLRQDQGWHPSRUDORUJDQL]DWLRQRIWURSLFDODQGWHPSHUDWHVSHFLHV DVVHPEODJHVRIORWLF&KLURQRPLGDHJournal of the Kansas Entomological Society 71: 388- 

CONNELL, J.H. :  'LYHUVLW\LQWURSLFDOUDLQIRUHVWVDQGFRUDOUHHIVScience 199:1302–1310.

'DNRWD&RXQW\6RLODQG:DWHU&RQVHUYDWLRQ'LVWULFW  &KXE&UHHNZDWHUVKHGDVVHVVPHQW5HSRUWIRUWKH1RUWK&DQQRQ5LYHU:DWHUVKHG0DQDJHPHQW 2UJDQL]DWLRQSSKWWSZZZGDNRWDFRXQW\VZFGRUJZDWHUVKHGVZDWHUTXDOLW\FKXE KWP

FERRINGTON, L.C., JR., M.A. BLACKWOOD, C.A. WRIGHT, N.H. CRISP, J.L. KAVANAUGH, & F.J. SCHMIDT:  $SURWRFROIRUXVLQJVXUIDFHÀRDWLQJSXSDOH[XYLDHRI&KLURQRPLGDHIRUUDSLGELRDVVHVVPHQW RIFKDQJLQJZDWHUTXDOLW\,QSediment and stream water quality in a changing environment: trends and explanations. HGV1(3HWHUV '(:DOOLQJ SS,$+63UHVV 2[IRUGVKLUH8.

HINTZE, J:  1&66DQG3$661XPEHU&UXQFKLQJDQG6WDWLVWLFDO6\VWHPV.D\VYLOOH87

KEIPER, J.B., & B.A. FOOT:  %LRORJ\DQG/DUYDO)HHGLQJ+DELWVRI&RH[LVWLQJ+\GURSWLOLGDH 7ULFKRSWHUD IURPD6PDOO :RRGODQG6WUHDPLQ1RUWKHDVWHUQ2KLR$nnals of the Entomological Society of America 93

REICE, S.R.  1RQHTXLOLEULXPGHWHUPLQDQWVRIELRORJLFDOFRPPXQLW\VWUXFWXUHAmerican Scientist 82 

VANNOTE, R.L. & B.W. SWEENEY:  *HRJUDSKLFDQDO\VLVRIWKHUPDOHTXLOLEULDDFRQFHSWXDOPRGHOIRUHYDOXDWLRQWKHHIIHFWRI QDWXUDODQGPRGL¿HGWKHUPDOUHJLPHVRQDTXDWLFLQVHFWFRPPXQLWLHVThe American Naturalist 115

'DWHUHFHLYHG 2008 Proceedings of the 16th International Chironomid Symposium 199

EMERGENCE OF ABLABESMYIA MONILIS (DIPTERA: CHIRONOMIDAE) FROM FINDLEY LAKE DURING WARM AND COLD YEARS

By TRUMAN E. SHERK1* & GREG H. RAU2

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ABSTRACT. 7KHUHZDVFRQVLGHUDEOH\HDUO\YDULDWLRQLQWKHPD[LPXPVXUIDFH WHPSHUDWXUHDW)LQGOH\/DNHDQGWKHGHSWKWRZKLFKWKHZDUPVXUIDFHZDWHUPL[HG GXULQJWKHVXPPHUAblabesmyia monilisHPHUJHGZKHUHWKHZDWHUWHPSHUDWXUHZDV PRUHWKDQž&7KHPD[LPXPHPHUJHQFHZDVIURPWKHRUJDQLFGHWULWXVIURPWKH VXUURXQGLQJIRUHVWDWDGHSWKRIPHWUHVLQ\HDUVZKHQWKHZDWHUZDVZDUPHQRXJK DWWKDWGHSWK

RESUMO. (QFRQWUDPRVXPDYDULDomRFRQVLGHUDYHOQDWHPSHUDWXUHPi[LPDQD VXSHU¿FLHGR/DJR)LQGOH\EHPFRPRQDSURIXQGLGDGHGDFLUFXODomRGDiJXDTXHQWH QRVPHVHVYHUQDLV$HFORVmRGHAblabesmyia monilisRFFRUHXTXDQGRDWHPSHUDWXUD GDiJXDH[FHGHXžFHQWtJUDGRV(FORVmRPi[LPDRFRUUHXQRVGHWULWRVRUJkQLFRV GHULYDGRVGDÀRUHVWDFLUFXQGDQWHQXPDSURIXQGLGDGHGHPHWURVTXDQGRDiJXD QDTXHODSURIXQGLGDGHDWLQJLXXPDWHPSHUDWXUDVX¿FLHQWPHQWHTXHQWH

*1&RUUHVSRQGLQJDXWKRU32%R[%UDQIRUG&786$7OWVKHUN#JPDLO FRP 26DQG\5RDG&DVWUR9DOOH\&$86$7OJKUDX#VEFJOREDOQHW

Bol. Mus. Mun. Funchal, Sup. N.º 13: 199-205, 2008 ISSN 0870-3876 200 Boletim do Museu Municipal do Funchal (História Natural) Sup. No. 13

INTRODUCTION

Species of Chironomidae emerging from Findley Lake in the warm year of 1973 emerged from depths between 0 m and 19.3 m, or 0.5 m and 19.3 m, except Ablabesmyia monilis (LINNAEUS, 1758), which was restricted to the shallow littoral (0 m to 5.6 m depth) (SHERK & RAU, 1992; 1996), indicating a higher temperature threshold compared to other species. Temperature threshold for for A. monilis emergence in the Pyrenees was estimated at 16° C to 17° C (LAVILLE, 1971). Annual maximum surface temperature at Findley Lake varies due to the variation in snowfall and therefore onset of the thaw (HENDREY & WELCH, 1974; SHERK & RAU, 1996). In some years maximum surface temperature exceeded 16° C, but in other years it was less than 16° C. Earlier papers (SHERK & RAU, 1992; 1996) have only explored emergence of A. monilis in 1973 from Findley Lake when maximum surface temperature reached 19.25° C. In this paper we study yearly variation in A. monilis emergence during years when the maximum surface temperature was above or below the emergence threshold of 16° C.

Study area Findley Lake (1128 m asl) in the Cascade Mountains of Washington, USA is surrounded by coniferous forest, talus slopes and wet meadows (Fig. 1). The glacial lake, carved in volcanic rock (andesite), has a maximum depth of 27.5 m, a mean depth of 7.8 m and an area of 11.4 ha. The lake is oligotrophic (HENDREY & WELCH, 1974). Most phytoplankton production is between the depths of 5 to 15 m, with the maximum production at 15 m. Detritus from the forest enters the lake near the shore (RAU, 1976), especially near the temporary melt-water tributaries.

Fig. 1. Map of Findley Lake with depths in metres. Locations of traps where Ablabesmyia monilis emerged (solid squares). Locations of traps where A. monilis did not emerge (hollow squares). Maximum depths of the east and west basins (triangles). 2008 Proceedings of the 16th International Chironomid Symposium 201

MATERIALS & METHODS

Insects were collected in 1.0 m2 ÀRDWLQJ emerJHQFH WUDSV covered by 1.0 mm mesh ¿EHrJODVV VFUHHQLQJ (SHERK et al., 2003). 6LPLODU WUDSV were SODFHG with one side onshore. Ten ÀRDWLQJ DQG seven shore WUDSV were used in 1972. Fewer WUDSV were used in other \HDUV Collections were PDGH DW one to ¿YH GD\ LQWHUYDOV GXULQJ ice-free SHULRGV

RESULTS

There ZDV FRQVLGHUDEOH \HDUO\ YDULDWLRQ in PD[LPXP VXUIDFH WHPSHUDWXUH DQG the GHSWK to which the ZDUP VXUIDFH ZDWHU PL[HG GXULQJ the summer )LJ 2). A. monilis VWDUWHG to emerJH from most GHSWKV when WHPSHUDWXUH UHDFKHG D threshold of 16° C. 0D[LPXP emerJHQFH ZDV from orJDQLF detritus derived from the VXUURXQGLQJ forest DWDGHSWKRIPLQ\HDUVZKHQWKHZDWHUZDVZDUPHQRXJKDWWKDWGHSWK In 1972 ODNH WKDZ 7 July), PD[LPXP VXUIDFH WHPSHUDWXUH UHDFKHG 21.0° C, but cold WHPSHUDWXUHV UHPDLQHG below the VXUIDFH )LJ 2). A. monilis only emerJHG within one metre of shore )LJ 3) when the VXUIDFH WHPSHUDWXUH UHDFKHG 16° C DQG FRQWLQXHGDWDZHWPHDGRZZKHQWKHVXUIDFHWHPSHUDWXUHIHOOEHORZƒ&

)LJ 2. THPSHUDWXUHV DW 0 m, 5 m DQG 10 m GHSWKV DW Findley /DNH from 1972 to 1974 DQG DW 0 m in 1975. THPSHUDWXUHV DW the shoreline of the wet PHDGRZV were sometimes ZDUPHU WKDQ where WKHYHUWLFDOWHPSHUDWXUHSUR¿OHZDVPHDVXUHG 202 Boletim do Museu Municipal do Funchal (História Natural) Sup. No. 13

In 1973 (early thaw on 7 June), maximum surface temperature reached 19.25° C, with warmer temperatures below the surface compared to other years (Fig.2), with the temperature EULHÀ\ reaching 18° C at 5 m depth. A. monilis emerged from the 0 to 5.6 m depth (Fig. 3 and SHERK & RAU, 1996), except at the 2.7 m deep site where there was less forest detritus (RAU, 1976). Emergence started when the bottom temperature reached approximately 16° C, (excluding the two individuals that emerged early at the 1.8 and 5.6 m deep sites near the tributaries) In 1974 (late thaw on 31 July), maximum surface temperature was only 12.5° C (Fig. 2). Only one individual emerged from the 1.8 m deep detritus near a small tributary (Fig. 3). In 1975 (thaw 7 July), maximum surface temperature was 17.4° C in mid August (Fig. 2). A. monilis emerged from the warm 0.5 m and 1.8 m deep sites where there was most allochthonous material (Fig. 3).

Fig. 3. Number of Ablabesmyia monilis that emerged per square metre per day at Findley Lake from 1972 to 1975. 2008 Proceedings of the 16th International Chironomid Symposium 203

 0D[LPXPHPHUJHQFHRIA. monilisZDVLQIURPWKHPGHHSVLWH ZLWKKLJKOHYHOVRIDOORFKWKRQRXVPDWHULDOIURPWKHVXUURXQGLQJIRUHVW 5$8  ([FHSWIRUDIHZLQGLYLGXDOVIURPWKHDQGPGHHSVLWHVLQQRA. monilis HPHUJHGIURPWKHWRPGHSWKVZKHUHSULPDU\SURGXFWLRQSUHGRPLQDWHG +(1'5(< :(/&+ 

DISCUSSION

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'DWHUHFHLYHG ! 2008 Proceedings of the 16th International Chironomid Symposium 207

CHIRONOMUS OCEANICUS PACKARD, 1869, A NEW JUNIOR SYNONYM OF HALOCLADIUS VARIABILIS (STAEGER, 1839) (DIPTERA: CHIRONOMIDAE)

By BROUGHTON A. CALDWELL1

With 1 table

ABSTRACT: A. S. Packard, Jr. described Chironomus oceanicus in 1869 based upon adult and immature specimens collected from the harbor at Salem, Massachusetts, USA. It has since been infrequently reported in the literature, and usually not based upon new material. The species has been most recently recognized as Cricotopus oceanicus (Packard). Some recently collected adult and pupal specimens from the 0DVVDFKXVHWWVFRDVWOLQHWHQWDWLYHO\LGHQWL¿DEOHDVWKLVVSHFLHVSURPSWHGIXUWKHU study to resolve their identity. Some questions as to the actual date of description and source, as well as possible location of type specimens were addressed. Additional adult specimens were examined from Massachusetts, as well as a very similar species documented from Maine, Canada, Finland and Norway locations to assess possible synonymy and variation. Close morphological similarity as well as distribution, ecology and life history details suggest interpreting Chironomus oceanicus as a junior synonym of Halocladius variabilis (Staeger).

Keywords: Chironomidae, Chironomus oceanicus, Cricotopus oceanicus, Halocladius variabilis, marine, synonym

RESUMO: A.S. Packard, Jr. descreveu Chironomus oceanicus em 1869, utilizando espécimes adultos e recolhidos no porto de Salem, Massachusetts, EUA, como referência. Desde essa data a espécie tem sido relatada de forma esporádica na literatura. No entanto, estas referências habitualmente não se baseavam em novo

1 2645 Muskogee Lane, Braselton, Georgia, 30517 USA, and Research Associate, Florida State Collection of , Gainesville, Florida, 32614 USA. E-mail: [email protected]

Bol. Mus. Mun. Funchal, Sup. N.º 13: 207-212, 2008 ISSN 0870-3876 208 Boletim do Museu Municipal do Funchal (História Natural) Sup. No. 13

material biológico. A espécie foi posteriormente reconhecida como sendo Cricotopus oceanicus (Packard). Espécimes adultos e pupas recolhidos recentemente na linha FRVWHLUDGH0DVVDFKXVHWWVHLGHQWL¿FDGRVFRPRSHUWHQFHQGRDHVWDHVSpFLHSHUPLWHP agora o desenvolvimento de novos estudos para resolver o problema de identidade taxonómica. Diversos problemas relacionados, com as datas de descrição, origem e localização dos espécimes tipo serão analisados. Espécimes adultos adicionais originários de Massachusetts, do Maine, Canada, Finlândia e Noruega serão estudados, por forma a determinar se estamos perante sinonímias ou variações. A proximidade morfológica, a distribuição, a ecologia e o ciclo de biológico sugerem que Chironomus oceanicus deve ser considerado como um sinónimo juvenil de Halocladius variabilis (Staeger).

A phenomenal naturalist, A. S. Packard, Jr. described two species of chironomid midges during his distinguished career. One of these, Chironomus oceanicus was described from Salem Harbor, Massachusetts (PACKARD 1869a). Prior brief accounts as Chironomus sp. were presented in meetings of the Essex Institute and Boston Society RI1DWXUDO+LVWRU\DQGUHÀHFWHGLQSXEOLVKHGPHHWLQJPLQXWHVDVZDVDVDQLOOXVWUDWHG description, but none of these established a valid name. A descriptive account and valid name appeared in an entomology textbook originally available in ten parts from the author (PACKARD 1869b). The Diptera section was published in March 1869 as was PACKARD (1869a). With no more precise publication dates available, I resolve this case RIVLPXOWDQHRXVSXEOLFDWLRQDFWLQJDV)LUVW5HYLVHU¿[LQJWKHSUHFHGHQFHRIWKHQDPH Chironomus oceanicus PACKARD (1869a) over the same name in PACKARD (1869b). Packard also informally characterized the species, at one time mistaking it for the of a rove beetle (Micralymna – later as Micralymma) living in green seaweed at the low water mark in Casco Bay, Maine. There are additional Massachusetts records of C. oceanicus from studies in Vineyard Sound, Buzzard’s Bay, Martha’s Vineyard, and adjacent waters reported by VERRILL (1873). The species was noted from the rocky shores of sounds and bays as well as on eel grass, oyster beds, and on pilings of wharves. JOHANNSEN (1905) reported on adults of what he thought to be this species from Woods Hole, Massachusetts, treating them in his key as Orthocladius oceanicus and above the description section as Orthocladius (?) oceanicus. JOHANNSEN (1952) placed the species in Cricotopus and provided a key for recognition of the female. The species has been included in several catalogs of North American chironomids, but reports have not been based on any new material for over a century. SUBLETTE (1967) examined three (one male and two females) of eight pinned specimens treated by JOHANNSEN (1905) from Woods Hole, noting that only the female was “adequately described” and that the species should be recognizable from the key by JOHANNSEN (1952). Females were considered recognizable among other features by a 2008 Proceedings of the 16th International Chironomid Symposium 209

dull thorax with contrasting vittae, thoracic setation, hairy eyes, genitalia without elongate FHUFLDQGOHJVQRWEDQGHG)RUWKHSUHVHQWVWXG\¿YHRWKHUSLQQHG EXWYHU\IUDJPHQWHG  individuals of Johannsen’s series were examined, and all (8) slide mounted. These as well as all other specimens examined in this study had morphology, measurements, and ratios as well as long looped seminal ducts, similar to those described for H. variabilis (HIRVENOJA 1973). One of the Johannsen females (as well as 2 Newfoundland females) exhibited what may be a very small seta on one antennal scape, but these could also be artifacts or debris. It has been surmised that Chironomus oceanicus might be a synonym of Chironomus variabilis Staeger (1839). HIRVENOJA (1973) in his classic work on the genus Cricotopus erected the genus Halocladius to include variabilis and other salt water species. He stated that C. oceanicus Packard had often been mentioned as a synonym of Trichocladius vitripennis (Meigen, 1818) in the sense of Thienemann, which included both H. variabilis and H. varians according to HIRVENOJA (1973). However, the true Chironomus vitripennis Meigen remains a nomen dubium as M. Spies (pers. comm.) has found the Meigen material in Paris to contain mixed species, none of which belong to Halocladius. Both Chironomus oceanicus and H. variabilis were described from marine coastal environments. In establishing Halocladius, HIRVENOJA (1973) recorded H. variabilis (under various names in the literature) distribution over parts of coastal Europe and Scandinavia, parts of the Soviet Union, Greenland, and Canada. Some more recent accounts (see below) that also include some aspects of ecology and phenology for North America include salt marshes and intertidal rocky coastline of the northeastern Atlantic coast of Canada. Chironomus oceanicus is known only from Massachusetts with no new material reported there since JOHANNSEN (1905). Massachusetts specimens recently collected by the author helped to prompt further study of the status of C. oceanicus. GARBURY et al. (2005) described a unique symbiosis between H. variabilis and the brown alga, Elachista fucicola, an epiphyte on the intertidal brown alga, Ascophyllum nodosum. Extremely high larval midge densities were reported and this association obviously represents a favored habitat. Such symbiosis may be restricted to the western Atlantic around Nova Scotia, but this association is certainly not obligatory for the midge. COLBO (1996) reported H. variabilis from intertidal algae, particularly Pilayella littoralis on the Newfoundland coast and GIBERSON et al. (2001) added a record from Prince Edward Island salt marshes, a habitat in which the occurrence of Elachista is unlikely. Different plant preferences were observed by PACKARD (1869a) in his description of Chironomus oceanicus larvae from an eelgrass (Zostera marina, commonly known as VHDZUDFN DQGDOVRIURPJUHHQ¿ODPHQWRXVVHDZHHGV From the similarities of morphology, measurements and ratios, the similar ecology, often with a preference for marine algae and seaweeds, and the overlapping distribution in the Nearctic, the present author concludes that Chironomus oceanicus 210 Boletim do Museu Municipal do Funchal (História Natural) Sup. No. 13

Packard (= Cricotopus oceanicus (Packard sensu Johannsen 1952 and Sublette 1967)) is a junior synonym of Halocladius variabilis (Staeger). Chironomus oceanicus has been a largely unreported taxon for over a century, probably due to lack of types and adequate descriptions. The few instances in which variation observed for Chironomus oceanicus falls outside that previously reported for H. variabilis are minor and are thus FRQVLGHUHGDVLQWUDVSHFL¿FYDULDWLRQ(YHU\VSHFLPHQH[DPLQHGLQWKLVVWXG\IURPWKH area of northeastern North America is clearly assignable to H. variabilis and there is no doubt that Chironomus oceanicus is a junior synonym. The Woods Hole specimens examined by SUBLETTE (1967) compare favorably to data given for H. variabilis by HIRVENOJA (1973). Measurements reported by

SUBLETTE (1967) for 2 females included a range of LR1 of 0.64-0.68, LR2 0.41, LR3 of 0.58, wing length 2.26-2.44 mm, and VR 1.11-1.13. The wing of 2.44 mm when measured from arculus to tip was 2.12 mm. The other slide mounted female wings were not accurately measurable. For the male, the LR2 was 0.40, LR 3 was 0.55, wing length was 2.00 mm (arculus to tip 1.71 mm), and VR was 1.12. Based on general features and collection location, a recently collected female from Massachusetts could be considered to represent Crictopus oceanicus (Packard). Meristic data are as follows in Table 1 and compare favorably with data of HIRVENOJA (1973) for H. variabilis. Other data include wing length of 2.13 mm, 24 squamal setae,

R setae 18, R4+5 setae 18, brachiolum setae 3, and VR 1.15. 7KHODUYDHDYDLODEOHIURP&DQDGDIRUWKHSUHVHQWVWXG\ZHUHLGHQWL¿DEOHDVH. variabilis. The average AR of the specimens measured (n=12) was 1.27 with a range of 1.15-1.36. All pupae and pupal exuviae examined in this study, including exuviae from Massachusetts, conformed to the description for H. variabilis.

7$%/(/HJPHDVXUHPHQWVDQGUDWLRV0DVVDFKXVHWWVDGXOWIHPDOHLGHQWL¿DEOHDVCricotopus oceanicus (Packard).

Fe Ti Ta1 Ta 2 Ta 3 Ta 4 Ta 5 LR SV BV p1 823 979 623 322 256 189 145 0.64 2.89 2.66 p2 878 901 367 211 178 33 122 0.41 4.85 3.33 p3 945 1056 600 334 278 167 145 0.57 3.33 2.82

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Massachusetts, Cornell University, and Peabody Essex Museum yielded no specimens. It is most likely that specimens were deposited at Yale College (now Yale University) but none could be located in the entomology or marine collections. Designation of a neotype at this time appears unnecessary due to synonymy. Material examined: Canada - Prince Edward Island, 29 June1993, leg. D. Giberson, 3 males; same data except 1 male, 2 females, 8 June1993; same data except 9 July 1993, 1 female; Manitoba, Fort Churchill, 1 August 1952, leg. J. G. Chillcott, 2 males, 1 female; Newfoundland, Maddox Cove near St. John’s, intertidal algae, 16 August 1993, leg. M. Colbo, 27 L; same data except 26 August 1993, 8 L, 1 Lex, 4 pharate male P, 2 pharate female P; same data except, sweep, intertidal rocks, 6 September 1993, 4 females; Broad Cove, near St. Phillip’s, intertidal algae, 1 June 1992, leg. M. Colbo, 4 L; Finland – Tvärminne Biological Station, 1952, leg. E. Palmén, 11 slides of males and females (body parts variously mounted), 12 Pex; Norway – Kviturdvikpollen, 3 May 1965, leg. R. Koskinen, 2 males, 1 female, 2 P, 1 Pex; same data except 26 April 1965, leg. R. Koskinen, 2 L; United States – Maine, York Co., Isles of Shoals, central valley pool, 11 August 2000, leg. W. K. Reeves, 1 male; Appledore Is., Shoals Marine Lab. (marsh near sewage treatment area), 25 June 1992, leg. S. K. Burian, 1 male; Massachusetts, Essex Co., tidal pool on bedrock, Rockport, Massachusetts, 27 August 2001, leg. B. A. Caldwell, 4 Pex; same data except, shoreline area, 26 August 2001, 1 female; Woods Hole, 2 males, 6 females, no other data (O. A. Johannsen collection, Cornell Univ.).

ACKNOWLEDGEMENTS

I thank B. Bilyj, S. K. Burian, M. Colbo, J. H. Epler, D. J. Giberson, M. Hirvenoja, E. R. Hoebeke, A. S. Johnson, J. Levy, B. Normark, P. D. Perkins, W. K. Reeves, J. E. Sublette, S. Werle, J. Winchell, and the late Nancy Adams for assistance. I especially thank M. Spies for review of the manuscript and help with translation of German language references.

BIBLIOGRAPHY

COLBO, M.: 1996. Chironomidae from marine coastal environments near St. John’s, Newfoundland, Canada. Hydrobiologia 318: 117-122.

GARBURY, D. J., M. M. JAMIESON, S. J. FRASIER, C. A. FERGUSON & P. S. CRANSTON: 2005. Ascophyllum (Phaeophyceae) and its symbionts. IX. A novel symbiosis between Halocladius variabilis (Chironomidae, Insecta) and Elachista fucicola (Elachistaceae, Phaeophyceae) from marine rocky shores of Nova Scotia. Symbiosis 40: 61-68. 212 Boletim do Museu Municipal do Funchal (História Natural) Sup. No. 13

GIBERSON, D. M, B. BILYJ & N. BURGESS:  6SHFLHVGLYHUVLW\DQGHPHUJHQFHSDWWHUQVRI1HPDWRFHURXVÀLHV ,QVHFWD'LSWHUD IURPWKUHH coastal salt marshes in Prince Edward Island, Canada. Estuaries 24: 862-874.

HIRVENOJA, M.: 1973. Revision der Gattung Cricotopus van der Wulp und ihrer Verwandten (Diptera, Chironomidae). Annales Zoologici Fennici 10: 1-363.

JOHANNSEN, O. A.: 1905. Aquatic nematocerous Diptera II. New York State Museum Bulletin 86 (Entomology 23): 76-331.

JOHANNSEN, O. A.: 1952. Family Tendipedidae (=Chironomidae) except Tendipedini. Connecticut State Geology and Natural History Survey 80: 3-26.

PACKARD, A. S.: 1869a. On insects inhabiting salt water. Communications Essex Institute 6: 41-46. 1869b. *XLGHWRWKHVWXG\RILQVHFWVDQGDWUHDWLVHRQWKRVHLQMXULRXVDQGEHQH¿FLDOWRFURSVIRU the use of colleges, farm-schools, and agriculturists (1st edition). Essex Institute Press, 702 pp.

SUBLETTE, J. E.: 1967. Type specimens of Chironomidae (Diptera) in the Cornell University Collection. Journal of the Kansas Entomological Society 40: 477-564.

VERRILL, A. E.: 1873. Report on the invertebrate of Vineyard Sound and the adjacent waters, with an account of the physical characters of the region. Pp. 295-778 in: Baird, S. F., Report on the FRQGLWLRQRIWKHVHD¿VKHULHVRIWKHVRXWKFRDVWRI1HZ(QJODQGLQDQG. United States Commission of Fish and Fisheries, 852 pp.

Date received: 06-04-2008. 2008 Proceedings of the 16th International Chironomid Symposium 213

A STRUCTURAL HERMAPHRODITE MICROPSECTRA (DIPTERA: CHIRONOMIDAE)

By PETER H. LANGTON*

With 1 Figure

ABSTRACT: A pharate imago Micropsectra atrofasciata from the River Chew, England, possessing well-developed male and female genitalia is described.

RESUMO: Uma farada de Micropsectra atrofasciata do Rio Chew, Inglaterra, possuindo genitália bem desenvolvida de macho e fêmea é descrito.

INTRODUCTION

Chironomid material collected in a drift net from the River Chew, Somerset, south west England, by R.S.Wilson on 25 April 1973 contained a pharate imago Micropsectra atrofasciata (Kieffer) with unusual genitalia.

METHOD

The specimen was heated in strong potassium hydroxide solution at 1000C for ¿YHPLQXWHVWRUHPRYHLQWHUQDOWLVVXHVQHXWUDOLVHGLQJODFLDODFHWLFDFLGGHK\GUDWHGLQ 100% isopropanol and mounted in Euparal dorsal surface uppermost.

RESULTS

The pupal armament of tergites III and IV determine the species to be Micropsectra atrofasciata (Kieffer 1911)(see LANGTON & VISSER 2003, STUR & EKREM 2006).

*University Museum of Zoology, Cambridge, Downing Street, Cambridge (address for correspondence: 5, Kylebeg Avenue, Coleraine, Northern Ireland, BT52 1JN, Tel. (+44) 2870328028, Fax. (+44) 2870328028, [email protected])

Bol. Mus. Mun. Funchal, Sup. N.º 13: 213-216, 2008 ISSN 0870-3876 214 Boletim do Museu Municipal do Funchal (História Natural) Sup. No. 13

The head and thorax are typically male (Fig. 1F), the antennae with a full complement RIÀDJHOORPHUHVDQGIRUHPHWDWDUVXVORQJ+RZHYHUWKHSRVWHULRUDEGRPHQKDVD vertical succession of male and female genital structures. The most dorsal are the female cerci (Fig.1A), followed by the male superior volsella and digitus (Fig.1B). Below these are the male median and inferior volsellae (Fig. 1C). Ventrally are the gonapophyses of segment VIII and vagina of the female genitalia (Fig. 1D). Situated further forward in the abdomen are the seminal capsules, notum and spermathecal ducts complete to the vaginal area (Fig. 1E).

DISCUSSION Hermaphroditism is almost unknown in insects apart from occasional developmental anomalies (NORMARK 2003). Partial hermaphrodites and gynandropmorphs have been reported for the Chironomidae (e.g. ANDERSEN & WANG 1997, SÆTHER & GALLOWAY EXWWKLVZRXOGDSSHDUWREHWKH¿UVWVSHFLPHQWREHUHSRUWHGZLWKD full complement of male and female genital structures. It is possible because the female appendages are derived from segment X (cerci) and ventrally on segment VIII (SÆTHEr 1977) while those of the male originate mainly from segment IX (SÆTHER 1980). Many abnormalities of development of the sexual apparatus of Chironomidae are induced by mermithid parasitism, but no evidence of this is detected in this specimen. Because the specimen was macerated before mounting it is not possible to determine whether it was a true hermaphrodite bearing both ovaries and testes.

ACKNOWLEDGEMENTS

I am grateful to Dr. R.S. Wilson for providing me with this specimen.

BIBLIOGRAPHY

ANDERSEN, T. & WANG, X.: 1997. Darkwinged Heleniella Gowin, 1943 from Thailand and China (Insecta, Diptera, Chironomidae, Orthocladiinae). Spixiana 20: 151-160.

LANGTON, P. H. & VISSER, H: 2003. Chironomidae exuviae. A key to pupal exuviae of the West Palaearctic Region. Amsterdam: Biodiversity Center of ETI (CD-ROM). 2008 Proceedings of the 16th International Chironomid Symposium 215

A B

C D

E F

Fig. 1. Micropsectra atrofasciata hermaphrodite A. cerci, B. superior volsella and digitus, C. median and inferior volsellae, D. gonapophysis VIII and vagina, E. cephalothorax, F. posterior abdomen, general view. (scale line = 0.1mm) 216 Boletim do Museu Municipal do Funchal (História Natural) Sup. No. 13

NORMARK, B. B.: 2003. The Evolution of Alternative Genetic Systems in Insects. Annual Review of Entomology, 48: 397-423.

SÆTHER, O. A.: 1977. Female genitalia in Chironomidae and other : morphology, phylogenies, keys. Bulletin of the Fisheries Research Board of Canada 197: 1-209. 1980. Glossary of chironomid morphology terminology (Diptera: Chironomidae). Entomologica Scandinavica, Supplement 14: 1-51.

SÆTHER, O. A. & GALLOWAY, T. D.: 1980. Sexual anomalies in Chironomini (Chironomidae: Diptera) from Lake Winnipeg, Manitoba, with observations on mermithid (Nematoda) parasites. Acta Universitatis Carolinae – Biologica, 1978: 193-211.

STUR, E. & EKREM, T.: 2006. A revision of West Palaearctic species of the Micropsectra atrofasciata species group (Diptera: Chironomidae). Zoological Journal of the Linnaean Society, 146:165-225.

Date received: 06-04-2008. 2008 Proceedings of the 16th International Chironomid Symposium 217

A NEW SPECIES OF PELOMUS REISS, 1989 (DIPTERA: CHIRONOMIDAE) FROM SOUTHEASTERN BRAZIL, WITH THE DESCRIPTION OF IMMATURE STAGES

By S. TRIVINHO-STRIXINO* & G. STRIXINO

With 12 Figures

ABSTRACT: The male, female, pupae and 4th instar larvae of a new species of the Harnischia-complex, from southeastern of Brazil, Pelomus psammophilus sp.n. DUHGHVFULEHGDQG¿JXUHG7KHODUYDHUHDUHGLQWKHODERUDWRU\WRREWDLQDOOOLIHVWDJHV were collected on littoral sand bottom of a little reservoir in central area of the state of São Paulo, Brazil.

RESUMO: As formas adultas e os imaturos de uma nova espécie do complexo Harnischia, da região sudeste do Brasil, Pelomus psammophilus sp.n. são descritas e ilustradas. As larvas, criadas no laboratório para obtenção de todos os estágios de vida, foram coletadas no sedimento arenoso de um pequeno reservatório localizado na área central do Estado de São Paulo, Brasil.

Keywords: Chironominae, Harnischia-complex, neotropics

*Corresponding author.1 Laboratório Entomologia Aquática / Departamento de Hidrobiologia / Universidade Federal de São Carlos, São Carlos, SP, Brasil. Cx. Postal: 676, CEP: 13.565-905. E-mail: [email protected], [email protected]

Bol. Mus. Mun. Funchal, Sup. N.º 13: 217-225, 2008 ISSN 0870-3876 218 Boletim do Museu Municipal do Funchal (História Natural) Sup. No. 13

INTRODUCTION

Pelomus, a genus of Harnischia-complex was erected by REISS (1989) based on specimens from Amazon region (Brazil). The hypopygium of P. notabilis, the only species whose adult is known, presents unusual features of the Harnischia complex. Nevertheless, the characteristics of its pupa, together with the other two described species (P. secundus Reiss, 1989 and P. tertius5HLVV MXVWL¿HGWKHLQFOXVLRQRIWKLVJHQXVZLWKLQWKH HarnischiaFRPSOH[%RWKWKHODUYDDQGZLWKVRPHPRGL¿FDWLRQWKHSXSDRIPelomus will key to Paracladopelma Harnisch which includes Saetheria Jackson in SÆTHER (1977a). Saetheria (JACKSON 1977) appears to be the closest related genus. The occurrence of unnamed species of this complex living on sand sediments of DUWL¿FLDOSRQGVLQWKHVWDWHRI6mR3DXOR6RXWKHDVWHUQ%UD]LOZDVUHFRUGHGE\675,;,12 75,9,1+2675,;,12%DVHGRQWKHPDWHULDOIURPWKLVSUHYLRXVVWXG\DQGRQ reared larvae and associated pupal exuviae and adults, we here describe a new species of PelomusLQFOXGLQJWKH¿UVWGHVFULSWLRQRIWKHODUYDOVWDJHIRUWKHJHQXV

MATERIAL AND METHODS

The material examined was mounted on slides in Euparal® or Hoyer’s medium. The general terminology follows SÆTHER (1977b, 1980). The term ‘taeniae’ was used IRUWKHÀDWWHQHGVHWDHRQWKHSXSDHDEGRPHQLQDFFRUGDQFHZLWK/$1*721   Measurements are given as the values of the holotype (when applicable) followed by values of the smallest and largest paratype, respectively. Seta counts are given as the range only. The holotype and most paratypes are deposited in the Laboratório de Entomologia Aquática collection, Universidade Federal de São Carlos, São Paulo, Brazil (UFSCar). One male paratype will also be deposited in the Museu de Zoologia, Universidade de São Paulo, São Paulo, Brazil (MZUSP).

Pelomus psammophilus sp. n. (Figs. 1-12)

Harnischia (Cladopelma) nr. nais Townes sp1. Roback, 1960: 328 Harnischia (?) sp. 2 Trivinho-Strixino & Strixino, 1995:111

(W\PRORJ\IURP*UHHN³SVDPPR´ VDQG DQG³SKLOXV´ IULHQG LWUHIHUVWR sandy sediments in which the larvae inhabit. 2008 Proceedings of the 16th International Chironomid Symposium 219

Type material. Holotype: male, Brazil: São Paulo State, Salesópolis, Rio Claro, 3RoR9HUGH (VWDomR%LROyJLFDGH%RUDFpLD [/HJ&*)URHKOLFK

Paratypes: 1 male with pupa and larva exuviae, Brazil: São Paulo State, São Carlos, Fazzari reservoir (UFSCar Campus), 17. v. 2005, Leg. S.T. Strixino, MZUSP Coll. 1 female with pupa and larva exuviae, Brazil, São Paulo State, Pirassununga, CEPTA, 20. iii. 2002, Leg. L. C. Correia. 1 4th instar larva, Brazil, São Paulo State, São Carlos, Fazzari Reservoir (UFSCar Campus), 24.viii. 1987, Leg. L. Albuquerque. 1 4th instar larva, Brazil, São Paulo State, São Carlos, Mayaca pond (UFSCar Campus), 2. iv. 1988, Leg.S. T. Strixino. 1 4th instar larva, São Paulo State, São Carlos, Boa Vista Reservoir (Fazenda Canchim), 24. viii. 1987, Leg. L. Albuquerque.

Diagnosis. The male of P. psammophilus sp. n. approaches that of P. notabilis LQWKHJHQHUDOFRQ¿JXUDWLRQRIWKHK\SRS\JLXPGLIIHULQJE\WKHVKDSHRIWKHDQDOSRLQW more pointed in P. psammophilus, and by having the inferior volsella distally enlarged as in the unnamed PelomusRI5(,66 ¿J 7KHSXSDRIP. psammophilus differs from the other known species of the genus by the absence of spines on the posterolateral PDUJLQRIVHJPHQW9,,LWUHVHPEOHVP. secundus and P. tertius by the possession of scutal tubercles and P. secundus by the absence of the pair of spines on median posterior margin of sternite VIII. The larva of P. psammophilus resembles the larvae of the genera Harnischia Kieffer, Paracladopelma and SaetheriaGLIIHULQJIURPWKHWZR¿UVWE\WKHQXPEHURI antennal segments (6), and by the presence of premandibular brush and labral lamella.

DESCRIPTION

Adult male 7RWDOOHQJWKPP*HQHUDOFRORUDWLRQ\HOORZLVKEURZQ7KRUD[EURZQLVK with vitae and postnotum dark brown (Fig. 1). Abdomen yellowish brown with dark EURZQEDQGVRQGLVWDOPDUJLQRIWHUJLWHV,9, )LJ *HQLWDOLDOLJKW\HOORZLVK Head. Yellowish brown. Antenna with 11 segments, brownish, 1.16-1.24 mm ORQJ$5 0D[LOODU\SDOSV\HOORZLVKEURZQ3DOSRPHUHV  250 µm long. Clypeus with 14-15 setae. Temporals with 14 setae. 7KRUD[6FXWDOWXEHUFOHDEVHQW$FURVWLFKDOVGRUVRFHQWUDOVLQVLQJOHURZ SUHDODUVVFXWHOODUV 220 Boletim do Museu Municipal do Funchal (História Natural) Sup. No. 13

Figs. 1-3. Pelomus psammophilus, sp. n. male. 1. Thorax dorsal and lateral view. 2. Abdominal tergites. 3. Hypopygium, dorsal view left, ventral view right. (ScDOH¿J —P 

Wing. Length 1.82 mm, width 0.55 mm, transparent, with lightly pigmented veins; FCu slightly distal than RM; VR 1.24. R 2+3 ending proximal to R1. R, R1 and R4+5 setose. Squama with 10 long setae. Brachiolum with 2 setae. 2008 Proceedings of the 16th International Chironomid Symposium 221

Lengths (in µm) and proportions of legs:

Fe Ti Ta1 Ta2 Ta3 Ta4 Ta5 LR I 815 554 1185 569 446 369 154 2.14 II 723 723 446 230 169 77 61 0.62 III 861 830 600 307 261 138 108 0.72

+\SRS\JLXP )LJ $QDOSRLQW—PORQJDSH[VOLJKWO\ODUJHWKDQEDVH Superior volsella with broad setose base, bearing 4 long distal setae and strongly naked hook-like distal part. Inferior volsella long, distally larger bearing 7-8 long setae. *RQRVW\OXVQDUURZ—PORQJ*F*V 

Adult female. Total length 2.7 mm. Coloration as male. +HDG$QWHQQDOÀDJHOORPHUHV —P$5 3DOSRPHUHV  —P&O\SHXVZLWKVHWDH7HPSRUDOZLWKVHWDH Thoracic setal count: 2 acrostichals, 13 dorsocentrals, 6 prealars, 17 scutellars.

:LQJ/HQJWKPPZLGWKPP95 6TXDPDZLWKVHWDH

Leg segment lengths in µm and LR proportions:

Fe Ti Ta1 Ta2 Ta3 Ta4 Ta5 LR I 692 508 908 400 338 323 154 1.79 II 615 580 340 138 107 61 61 0.58 III 677 708 331 200 169 1107 92 0.46

*HQLWDOLD6WHUQLWH9,,,EHDULQJVHWDHLUUHJXODUO\GLVWULEXWHGRQHDFKVLGH *&D9,,,VWURQJURXQGHGFDXGRODWHUDOO\*S9,,,VOLJKWO\GLYLGHGLQWRGRUVRPHVDOOREH YHQWURODWHUDOOREHUHGXFHG3RVWJHQLWDOSODWHZHDN*RQRFR[LWH,; *F,; ZLWKVHWDH 6HJPHQW;ZLWKVHWDRQHDFKVLGH&HUFLUHODWLYHO\ODUJHVRPHZKDWUHFWDQJXODU µm long. Labia with weak microtrichia. Seminal capsule and spermathecal ducts not discernible in the slide preparation.

Pupa ([XYLDHSDOHEURZQ$EGRPHQPP PDOH ORQJPP IHPDOH  222 Boletim do Museu Municipal do Funchal (História Natural) Sup. No. 13

&HSKDOLFWXEHUFOHVVKRUWFRQLFDODSLFDOO\SRLQWHGIURQWDOVHWDHVKRUWLQVHUWHG subapically. Thorax (Fig. 4). Basal ring oval, somewhat constricted medially. Dorsum of thorax granulose anteriorly. Prealar tubercle rounded. Scutal tubercle prominent. Two precorneals, 2 antepronotals and 4 dorsocentrals closed spaced present. Abdomen (Fig. 5). Hook row, interrupted medially, occupying about 1/3 the width of segment II. Segment I with large anterior pedes spurii B. Pedes spurii B absent RQVHJPHQW,,3HGHVVSXULL$SUHVHQWRQVWHUQLWH,97HUJLWHV,DQG,,EDUH,,,9ZLWK SRVWHULRUJURXSRIPHGLDQ¿HOGVRIVSLQHV9,,ZLWKWULDQJXODUPHGLDQ¿HOGRIODUJHVSLQHV &RQMXQFWLYHVEDUH3OHXUDEDUHH[FHSWIRUVPDOOSDWFKHVRI¿QHVSLQHVRQSRVWHULRUFRUQHU of segments V-VII. Anal lobe well developed with complete fringe of about 45 taeniae and one dorsal semitaeniate seta on each side.

Abdominal setation: Segment tergite sternite lateral I6 21 II 4 6 3 III 4 6 3 IV 4 6 3 V4 64T VI 4 4 4T VII 4 6 4T VIII 0 2 5T

4th instar larvae Q   Total length 5.2 mm. +HDG:LGWKȝPOHQJWK—P$QWHQQDVHJPHQWHGZLWKEODGHDULVLQJLQ PLGRIVHFRQGVHJPHQWEDVDOVHJPHQW—P  ORQJHUWKDQÀDJHOOXPZLWKEDVDO ULQJRUJDQLQGLVWDOVHJPHQWVKRUWHUWKDQVHJPHQW6W\OHSUHVHQWRQVHJPHQW (Fig. 10). /DEUXP )LJ 6,¿QHVKRUWVHWDOLNH6,,ORQJVWURQJEODGHOLNH6,,,YHU\ VKRUW6,9DHORQJDWHGVHJPHQWHG6,9EVKRUWDULVLQJEHKLQG¿UVWVHJPHQWRI6,9D /DEUDOODPHOODSUHVHQWGLYLGHGLQWRSDUWV3HFWHQHSLSKDU\QJLVDVLQJOHGLVWDOO\WUL¿G VFDOH3UHPDQGLEOH )LJ ZLWKODUJHDSLFDOWHHWKDQGVPDOOHUWRRWKSUR[LPDOO\EUXVK present. Mandible (Fig. 9). Lacking dorsal tooth, with strong apical tooth and three small LQQHUWHHWKORZHVWLQQHUWRRWKSDOHU3HFWHQPDQGLEXODULVZLWKWZR¿ODPHQWV 2008 Proceedings of the 16th International Chironomid Symposium 223

Maxilla with 81µm (78-83) long maxillary palp (Fig.8). Mentum (Fig. 11) with four wide light median teeth in a convex arc; three darker teeth and 4-5 short paler lateral teeth on either side. Third lateral teeth separate of second by a coarse suture. Ventromental plate about as wide as mentum, course striated with anterior margin weakly crenulated. Abdomen with anal tubules 275 µm long, with a median constriction, shorter than posterior parapods (Fig. 12).

Figs. 4 - 12. Pelomus psammophilus, sp. n. pupa (4-5) and larva (6-12). 4. Thorax, lateral view. 5. Abdominal tergites. 6. Labrum. 7. Premandible. 8. Maxillary palp. 9. Mandible. 10. Antenna. 11. Mentum and ventromental plate. 12. Posterior abdominal segments. 6FDOH¿JV 6, 7, 8, 9, 11 = 100 µm; 4, 5, 12 = 500 µm). 224 Boletim do Museu Municipal do Funchal (História Natural) Sup. No. 13

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The larvae of P. psammophilus sp. n. are relatively common in small rural reservoirs in the middle of the state of São Paulo. These, among others that probably belong to the same genus (Harnischia (?) sp. 1, Harnischia " VS75,9,1+2675,;,12  675,;,12 OLYHRQVDQGVHGLPHQWVRIVPDOOUHVHUYRLUVRURQVDQGOLWWRUDORIODUJHU RQHV 675,;,12 75,9,1+2675,;,12  *XWFRQWHQWDQDO\VLVRIVRPHVSHFLPHQVSRLQWRXWWKHSUHVHQFHRIDQLPDOUHPDLQV (Oligochaetes bristles) together with detritus. The buccal apparatus with pointed mandibles and mentum with median unsclerotized teeth indicate that the larva is capable to ingest larger food such as other invertebrates or large algae (Diatoms and Desmids).

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We are indebted to Dr. Fabio de Oliveira Roque for important suggestions and discussions. We also thank to Dr. Ole Saether for the comments and corrections on the manuscript. This work was partially supported by Fundação para o Amparo à Pesquisa do Estado de São Paulo (FAPESP) within the BIOTA/ FAPESP – The Biodiversity Virtual Institute Program (www.biota.org.br).

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-$&.621*$ 1977. Nearctic and Palaearctic Paracladopelma Harnisch and Saetheria n.gen. (Diptera: Chironomidae). Journal of the Fisheries Research Board of Canada, 34: 1321-1359.

/$1*7213+  ,IQRW³¿ODPHQWV´WKHPZKDW"Chironomus, 6: 9.

REISS, F.: 1989. Pelomus gen. n., ein weiterer potamobionter vertreter des Harnischia complexes aus dem Amazonasbecken (Chironomidae, Diptera). Acta Biologica Debrecina Oecologica. Hungaric, 2: 305-314.

ROBACK, S. S.:  7KH&DWKHUZRRGIRXQGDWLRQ3HUXYLDQ$PD]RQH[SHGLWLRQ;,,±'LSWHUDZLWKVRPH observations on the salivary glands of Tendipedidae. Monographie of Academy of Natural Science of Philadelphia, 14: 305-375. 2008 Proceedings of the 16th International Chironomid Symposium 225

SÆTHER, O.A.: 1977a. Taxonomic studies on Chironomidae: Nanocladius, Pseudochironomus, and the Harnischia complex. Bulletin of the Fisheries Research Board of Canada, 196: 1-143. 1977b. Female genitalia in Chironomidae and other Nematocera: morphology, phylogeny, keys. Bulletin of the Fisheries Research Board of Canada, 197: 1-209.  *ORVVDU\RIFKLURQRPLGPRUSKRORJ\WHUPLQRORJ\ 'LSWHUD&KLURQRPLGDH Entomologica scandinavica Supplement, 14: 1-51.

675,;,12* 75,9,1+2675,;,126  3RYRDPHQWRVGH&KLURQRPLGDH 'LSWHUD HPODJRVDUWL¿FLDLV,Q1HVVLPLDQ-/&DUYDOKR A. L. (Org.). Ecologia de Insetos Aquáticos (série Oecologia Brasiliensis,. 5: 141-154.

75,9,1+2675,;,126 675,;,12*.: 1995. /DUYDVGH&KLURQRPLGDHGR(VWDGRGH6mR3DXOR*XLDGHLGHQWL¿FDomRHGLDJQRVHGRV gêneros6mR&DUORV33*(518)6&$5S,/SODWHV

Date received: 06-04-2008.