2003. The Journal of Arachnology 31:1±7 SUB-FOSSIL SPIDERS FROM HOLOCENE PEAT CORES Alan G. Scott: Department of Earth Sciences, University of Manchester, Manchester M13 9PL, UK. E-mail: [email protected] ABSTRACT. An attempt was made to recover identi®able spider sub-fossils from peat cores taken from a post-glacial basin mire in Cheshire, north-west England. Although the features normally required to identify specimens to species level in taxonomic keys were rarely preserved, carapace morphology and cheliceral dentition allowed unequivocal identi®cation to species level in many cases. Current lack of knowledge of the autecology of wetland spiders prevents any conclusion regarding the paleoecological conditions, but the technique could reveal insights into the post-glacial development of the spider faunal assemblage of mires. Keywords: Spiders, sub-fossils, peat, chelicerae The value of undisturbed peat deposits as copy of the mature female epigyna and male datable sediment is well recognized and the palps. These are not always well sclerotized pollen record has been widely used to study and would, in any case, require some expertise the post-glacial vegetational succession, cli- to recognize if distorted and out of context. matic change and the development of agricul- However, Coope (1965, 1986) suggested that tural activity during the last 10,000 years (the (in the case of beetles) fragments of cuticle Holocene epoch). This technique depends on may be identi®able if a well determined ref- the remarkable resistance of the exine of pol- erence collection is available. The taxonomic len grains to natural decay and the chemical usefulness (at the supra-speci®c level) of other digestions used in their recovery, together morphological characters of spiders, has been with the taxonomic value of their morphology. reviewed by Lehtinen (1978), and these in- It is also recognized that identi®able (al- clude eye patterns, cheliceral armature and leg though disarticulated) invertebrate remains structure. can be recovered from peat and other uncon- As a reference collection of peatland spi- solidated sediments to give useful palaeoecol- ders, appropriate to the study area, was avail- ogical information. This has been particularly able to the author, he decided to attempt the successful in the case of beetles which have a retrieval of identi®able spider sub-fossils from heavily sclerotized exoskeleton that is resis- a species-rich basin mire in Cheshire, Eng- tant to decay and bears surface details which land. enable identi®cation to species level (Coope METHODS 1965). Also many beetle species can be linked Site description.ÐSeveral samples were closely to habitats. taken from the basin mire at Wybunbury Moss Hofmann (1986) was able to retrieve the National Nature Reserve, Cheshire, England head capsules of Chironomus larvae by essen- (National Grid Reference: SJ697503), on a tially the same method used for pollen anal- transect described by Green & Pearson (1977) ysis. Chironomid larvae and their exuviae and for which a stratigraphy was available. would have been particularly numerous in This site is in an area of northwest England peat and the head capsules are well sclero- which has an oceanic climate and glacial drift tized, but as I had observed the rapid disartic- geology which have favored the development ulation and decomposition of spiders in pitfall of numerous basin and raised mires. traps when ¯ooding had diluted the preser- Core sampling.ÐThe total depth of peat at vative, it seemed unlikely that identi®able spi- the study site is about 5 m, representing ac- der remains would be preserved. cumulation over 5000 years. Sampling depths Keys for the identi®cation of spider species were 650±750 mm (ca. 500 yr BP), 1150± have required, among other things, micros- 1250 mm (ca. 1000 yr BP) and 1650±1750 1 2 THE JOURNAL OF ARACHNOLOGY mm (ca. 1400 yr BP), these being the bottom 100 mm of each sequential core. This chro- nology is derived from the peat accumulation rate suggested for this site by Green & Pear- son (1977). The samples were obtained by us- ing a `Russian pattern' peat sampler (Jowsey 1966; Aaby & Digerfeldt 1986), which pro- duces half-cylinders of peat with a diameter of about 2.5 cm and length of 50 cm (Fig. 1). Extension rods can allow sampling to more than 5 m if suf®cient manpower is available to insert and withdraw the sampler at this depth but, for this single-handed investigation, the maximum depth used was 1.75 m. The surface vegetation was cut away before cor- ing, to minimize the possibility of contami- nation with recent material. Aaby & Digerfeldt (1986) described the procedure as shown in the inset cross-sections of Fig. 1: inserting the tool with the cutting shell over the rib and then an anti-clockwise rotation to cut a half-cylinder of peat. Jowsey (1966), on the other hand, performed the re- verse action, a clockwise rotation of the cut- ting shell producing quarter-cylinders of peat on either side of the rib. The method of Aaby & Digerfeldt was used, because insertion with the rib exposed increased the dif®culty of pen- etrating the upper layer which contains tough roots of Eriophorum and Erica. Samples were returned to the laboratory in `zip-seal' plastic bags. Should ®ner stratigraphic resolution be required, then the intact cores can be laid in split plastic pipes and wrapped in cling-®lm. Extraction of arthropod remains followed a modi®ed scheme of Coope (1986). Half kilo- gram aliquots of the peat (larger amounts clogged the sieves) were dispersed in warm water and the coarser material removed by washing through a 2 mm mesh sieve, the ma- terial passing through being collected on a 250 mm mesh sieve of 20 cm diameter. The Figure 1.ÐThe `Russian pattern' peat corer (after drained, but still damp, product was trans- Aaby & Digerfeldt 1986). ferred to a large bowl and mixed thoroughly for several minutes with its own volume of paraf®n (kerosene). This can be done by rub- transferred, either by decanting or spooning ber-gloved hand or with a table fork or similar off, to another sieve of smaller mesh (I used implement. 53 mm) and most of the residual paraf®n can After decanting as much surplus paraf®n as be removed by rinsing with warm water con- possible (which can be reused), water is added taining detergent. to the mixture, with stirring, to resuspend the Coope (1986) then suggested a ®nal rinse sample and ®ll the container. After ca. 1 hour in alcohol, followed by microscopy of the ex- the sample will have divided into sedimented tracted material in alcohol. While this may and ¯oating fractions. The ¯oating fraction is have worked well with the organic silts that SCOTTÐSUB-FOSSIL SPIDERS FROM PEAT 3 Table 1.ÐSummary of sub-fossil retrieval. Level jective impression that the yield improved 1 samples are from 650±750 mm depth (ca. 500 yr with experience of the technique, especially at BP); Level 2 samples are from 1150±1250 mm the microscopy stage. Chelicerae, at low mag- depth (ca. 1000 yr BP) and Level 3 samples are ni®cation, could have been confused with from 1650±1750 mm depth (ca. 1400 yr BP). small leg segments, such as trochanters from other arthropod taxa, and ignored. There Level seems no reason to doubt that similar yields 12 3 of spider sub-fossils would be available Amount of peat processed 5 kg 0.5 kg 0.5 kg throughout the peat column. Identi®cation of sub-fossils.ÐSpider spe- Carapaces 59 5 cies that were identi®able from sub-fossils Sternums 22 1 1 Chelicerae 130 2 3 came from the following categories: linyphiid Male palps 4 1 males that had distinctive head modi®cations; Palpal tibiae 1 larger species from which male palps or fe- Epigyna 1 male epigynes survived; larger species with distinctive cheliceral dentition; larger species with distinctive carapace features such as eye he was studying, the ¯oating fraction from pattern, hair and bristle distribution, overall partially humi®ed peat samples was often size and shape or some combination of these. bulky and contained much vegetable material. Some of the more interesting sub-fossils It was found that resuspending the ¯oating obtained from the material collected at Wy- fraction in clean water, after the detergent bunbury Moss are shown in Figs. 2±19. It is rinse, caused most of the vegetable matter notable that three small linyphiid species were (sphagnum leaves) to sink. identi®ed (Figs. 2±4) that did not ®gure in a The new ¯oating fraction was transferred to recent survey of this site (Felton & Judd a petri dish without alcohol treatment for mi- 1997): Hypselistes jacksoni (O.P.-Cambridge croscopy. Items of interest could be picked 1902), Gonatium rubens (Blackwall 1833) from the surface of the water with ®ne forceps and Savignya frontata Blackwall 1833. The and transferred to alcohol. Best visibility of males of these species have characteristic car- the sub-fossils (many of which are transparent apace pro®les, and the ®rst is a mire indicator and pale in color) was obtained by using a species. Other linyphiid carapaces were iden- mixture of transmitted and incident illumina- ti®able as species present in the modern fauna tion. A mechanical stage with X and Y axis (Figs. 5 & 7). controls is useful for scanning the surface of The majority of the remaining carapaces the water in a systematic manner. were probably from female linyphiids. Al- though well preserved, and several seemed RESULTS conspeci®c, they could not be positively iden- The bulk of the recognizable invertebrate ti®ed from the reference collection (one is remains consisted of large numbers of oribatid shown in Fig.