Macrogastra Badia (C. Pfeiffer

Vol. 17(2): 53–62 MACROGASTRA BADIA (C. PFEIFFER, 1828) (GASTROPODA: PULMONATA: CLAUSILIIDAE) IN ZIELENIEC (BYSTRZYCKIE MTS, CENTRAL SUDETES) – ECOLOGY, CONSERVATION STATUS AND LIFE HISTORY – PRELIMINARY DATA TOMASZ K. MALTZ, BEATA M. POKRYSZKO Museum of Natural History, Wroc³aw University, Sienkiewicza 21, 50-335 Wroc³aw, Poland (e-mail: [email protected], [email protected]) ABSTRACT: Information on thedistribution on theAlpine M. badia in Poland dates from the 1960s and was not verified subsequently. A new locality was discovered in 2003 (Bystrzyckie Mts, Zieleniec near Duszniki-Zdrój); it forms a part of a group of isolated, Polish and Czech localities on the border of the species’ distribution range. In the discussed part of the range the species is threatened by habitat destruction and climatic changes. It is legally protected in Poland but preserving its populations requires habitat protection. The preferred habitat is herb-rich beech forest, and cool and humid climate is crucial for the species’ survival. The composition of the accompanying malacofauna varies among the sites which is probably associated with their origin. M. badia is oviparous; in May and June it produces batches of 1–3 eggs. The eggs are partly calcified, 1.39–1.61 in major and 1.32–1.45 mm in minor diameter. The incubation period is 16–19 days; the hatching is asynchronous; the juveniles reach adult size in 7–8 months. Some data on shell variation are provided; the number of apertural folds varies more widely than formerly believed. KEY WORDS: Clausiliidae, endangered species, Macrogastra badia, lifehistory, ecology INTRODUCTION Macrogastra badia (C. Pfeiffer, 1828), one of the German side), Orlické hory and Králický Snìznik most rare species in the malacofauna of Poland, is – (LOZEK 1955, 1956, 1964, BRABENEC 1973, HLAVÁÈ & next to Cochlodina costata and Charpentieria ornata – HORSÁK 2002). In Poland it was first found by WIK- one of three native clausiliids representing the Alpine TOR (1964) in a few localities in the Sto³owe, Orlickie (East Alpine) zoogeographical element (LOZEK 1955, and BystrzyckieMts and in thenevirons of Gnie 1956, 1964, WIKTOR 1964, 2004a, b, KLEMM 1969, woszów near Bystrzyca K³odzka. Both the Czech and KERNEY et al. 1983, RIEDEL 1988, HLAVÁÈ & HORSÁK thePolish localities form a group of thenorth-east -- 2002). Its main distribution rangeincludes thesouth - ernmost, isolated and relic occurrences of the snail ern part of Eastern Alps (LOZEK 1955, 1956, 1964, (WIKTOR 1964, 2004a, b, RIEDEL 1988) (Fig. 1). WIKTOR 1964, KERNEY et al. 1983, RIEDEL 1988, HLAVÁÈ & HORSÁK 2002), in the west reaching the 1964, 2004a, b, KERNEY e t a.l 1983, RIEDEL 1988, Brenner Pass and the Bavarian Allgäu region; in the HLAVÁÈ & HORSÁK 2002, MALTZ & SULIKOWSKA- M. badia is a forest-dweller (LOZEK 1964, WIKTOR south it comprises Styria, northern Carinthia, the Al- -DROZD 2008), inhabiting deciduous and mixed forpine part of north-eastern Italy (KLEMM 1969), and - ests at higher altitudes (above 700 m a.s.l.), where it across the Styrian Alps – northern Slovenia (BOLE & stays on logs, in litter, under bark and stones, and SLAPNIK 1997). In the Czech Republic M. badia was re- sometimes on shaded, moss-covered rocks. corded from Šumava [Bohemian Forest] (also on the 54 Tomasz K. Maltz, Beata M. Pokryszko Fig. 1. Distribution of Macrogastra badia: A – in Europe, B – in the Sto³owe, Orlickie& BystrzyckieMts: [ ¡] – sites reported by WIKTOR (1964), [l] – Czech localities (according to BRABENEC 1973) In Poland the species is legally protected (ROZPORZ¥DZENIE MINISTRA ŒRODOWISKA 2004), it is Thelack of information on thebiology of thespe - <ul style="display: flex;"><li style="flex:1">cies, and thefact that thefaunistic data areinsuffi </li><li style="flex:1">-</li></ul>red-listed (WIKTOR & RIEDEL 2002) and included in cient to provide a basis for an efficient protection theRed Data Book (W IKTOR 2004b) as critically en- strategy, constitute a serious problem. Finding its dangered (CR) because of its few and small popula- prosperous population encouraged us to undertake a tions threatened by random events and habitat de- more detaled study (field and laboratory observa- <ul style="display: flex;"><li style="flex:1">struction (POKRYSZKO & MALTZ 2007). In theCzech </li><li style="flex:1">tions), in hopethat theresults might contributeto de - </li></ul>Republic it is regarded as endangered (JUØIÈKOVÁ et signing an adequate protection strategy. al. 2001) because of the small number of localities and similar risk factors. STUDY AREA In 2003 we discovered a new, previously unknown Alps and found above850 m a.s.l. (Œ WIERKOSZ 2003). locality of M. badia. It is situated in Zieleniec near Thedominant treis bech, sycamoremapleand Duszniki-Zdrój (Bystrzyckie Mts, Central Sudetes) spruce being less abundant; rowan forms an admix(Fig. 2A). It is a forest fragment ca. 0.9 ha in area, on ture, the herb layer is rich, especially in tall montane the eastern slope of the border mountain range, at an perennials of the class Betulo-Adenostyletea (Fig. 2B–D). altitudeof 900–920 m a.s.l., very closeto thevillage, ca. 200 m from theroad crossing its cne tre (50°20’21”N, 16°22’52”E, UTM: WR 97). Numerous rotting logs (deciduous) and the natural vertical and horizontal structureof thetreestand in dicate its spontaneous development since at least 80 - Floristically, it is a herb-rich beech forest (Aceri- years (ŒWIERKOSZ 2003). -Fagetum) – a plant community characteristic of the Macrogastra badia (C. Pfeiffer, 1828) in Zieleniec 55 Fig. 2. Siteof Macrogastra badia in Zieleniec (A) and its habitat (B-D) (Photo B. M. POKRYSZKO) <ul style="display: flex;"><li style="flex:1">Thesubstratum is micaschist with intercalations of </li><li style="flex:1">tion period is ca. 170 days (days with temperature </li></ul>amphibolites, quartzites and crystalline limestones, >5°C), the number of days with temperature >10°C is on which acid brown soils and podzols develop ca. 110, and there is no climatic summer (days with <ul style="display: flex;"><li style="flex:1">(STUPNICKA 1989). </li><li style="flex:1">temperature >15°C). Winter lasts more than 18 </li></ul>Pluviothermically, the region of Zieleniec is in- weeks, the annual precipitation sum is 1,000–1,200 cluded in class VI d (cool and very humid climate) mm, thesnowfall period is ca. 190–200 days (K OSIBA (SCHMUCK 1959, after WALCZAK 1968). The vegeta- 1955, after WALCZAK 1968). METHODS Gastropods were sampled on two occasions: in cessed as described in CAMERON & POKRYSZKO June and October 2007. On each occasion collecting (2005). Live clausiliids, helicids and slugs were identiinvolved visual search during at least 2 hrs, consider- fied in the site and released, the remaining material <ul style="display: flex;"><li style="flex:1">ing all possiblemicrohabitats, and taking a littre </li><li style="flex:1">was identified in the laboratory and deposited in the </li></ul>sample of 10–15 l volume. The samples were then pro- collection of the Natural History Museum, Wroc³aw 56 Tomasz K. Maltz, Beata M. Pokryszko Fig. 3. Macrogastra badia. A – activeadults , B – shell shown from two sides, C – aperture with parietal lamella (1), folds (2) and lower lamella (3). Scale bar 1 mm (Photo T. K. MALTZ) University. The nomenclature and systematic arrange- Peking cabbage, carrot and champignons (food exment follow KERNEY e t a.l (1983) and WIKTOR changed every 2–3 days). High humidity was main(2004a), ecological and geographical groupings were tained in theboxes through sprinkling thetissuepaadopted after RIEDEL (1988). The species composi- per, moss and timber with water every 2–3 days. From <ul style="display: flex;"><li style="flex:1">tion in the locality was compared with species lists thebeginning of theculturetill June2008 thetem </li><li style="flex:1">-</li></ul>from sites with M. badia availablein theliterature perature was 16–17°C, in July and August ca. 20–22°C, (HLAVÁÈ & HORSÁK 2002, HLAVÁÈ 2003), using the and since September – again ca. 17°C. Theboxes Nei index (IN) to estimatethesimilarity (P OKRYSZKO were kept in natural lighting regime. <ul style="display: flex;"><li style="flex:1">& CAMERON 2005). </li><li style="flex:1">In July 2008, shells of 39 adults from the laboratory </li></ul>In October 2007, 39 individuals of M. badia were culture were measured; the measurements included brought to the laboratory: 22 adults and 17 juveniles shell height and width (H, W) and aperture height (8 with shells of 6–6.9 whorls, 5 with shells of 7–7.9 and width (h, w); whorls were counted (N), as well as whorls and 4 with shells of 8 whorls). The snails were the folds between the superior and inferior apertural kept in perspex boxes of 6 × 12 × 15 cm, lined with tis- lamellae (Fig. 3). Height/width ratios of the shell sue paper, with tussocks of moss and pieces of dead (H/W) and aperture (h/w) were calculated. Meatimber from the locality. Adults were placed in groups surements were taken with calibrated eye-piece to the of 4–5 in separate boxes, juveniles – in groups corre- nearest 0.01 mm. Major and minor diameter of eggs sponding to the number of whorls. The boxes were (a, b) were measured to the nearest 0.01 mm, and a/b controlled every two days, the tissue paper was ex- ratio was calculated. Growth rate of juveniles brought changed every week. Dolomite tables were provided from thefield and thoseobtained in thelaboratory <ul style="display: flex;"><li style="flex:1">as thesourceof calcium. Thesnails werefed lettuce, </li><li style="flex:1">was monitored. </li></ul>RESULTS The species composition of the snail community in companying fauna is composed of 25 species, includthe locality in Zieleniec is shown in Table 1. The ac- ing 3 clausiliids and 6 slugs. Ecologically, thecommun - Macrogastra badia (C. Pfeiffer, 1828) in Zieleniec 57 ity includes three groups: forest-dwellers (majority), spherical, sometimes slightly elongated (a/b: euryoecious species and hygrophiles (Fig. 4). Zoogeo- 1.00–1.13; mean 1.08) (Fig. 8). Calcium carbonate in graphically it is composed of European (14 species), the egg envelopes does not form distinct, separate Holarctic (6) and Palaearctic (2) species, and one crystals but a morecontinuous layer. Theincubation species of each of the remaining categories (Fig. 5). period ranged from 16 to 19 days (mean 17.2). <ul style="display: flex;"><li style="flex:1">Themean similarity I </li><li style="flex:1">with theŠumava malacocoe - Hatching was asynchronous and lasted 1–2 days. The </li></ul>Nnoses is 0.52 (0.41–0.68), and that between the Czech hatching success ranged from 0 to 100 (mean 53%). and Polish sites – 0.44. Only threof thesnails hatched in thelaboratory The shells from Zieleniec are spindle-shaped and reached ultimate size (16% of obtained eggs (19) and rather tumid (H/W: 3.56–4.18; mean 3.82), purple- 30% of juveniles (10)). -brown, shiny, with rectangular apertures of slightly rounded bottom corners (h/w: 1.08-1.39; mean 1.25) field in October 2007 depended on their size upon col(Table2). Thenumbre of whorls is ca. 10 (N: lection: individuals with shells of 8 whorls completed Reaching ultimate size by snails brought from the 9.25–10.5; mean 9.87). The arrangement of folds in their growth at the begining of January 2008 (in ca. 2.5 the aperture varies widely (Figs 6, 7). The set of three months), thoseof 7 whorls – at theend of February folds is typical (nearly 40% of the shells), shells with (ca. 4 months), thoseof 6 whorls – at theend of March <ul style="display: flex;"><li style="flex:1">2.5 (two completeand onerudimentary fold) or 2 </li><li style="flex:1">(ca. 5 months). Thegrowth of individuals hatched in </li></ul>folds are rather frequent (12.75% and 10.2%, respect- thelaboratory to adult sizetook 7–8 months (2 snails ively). One specimen in the series has 4 folds; in some hatched at half of May 2008 finished their growth at instances the folds are connected. Egg laying in the laboratory was observed from 16 of June2008 – at theend of January 2009). May till 20 June 2008. A total of 19 eggs were pro- The temperature had a significant effect on the half of December, the snail hatched at the beginning duced, in 8 batches (1–3 eggs, mean 2.38) (Table 3). condition of thesnails (at high, constant humidity Theeggs of M. badia are resilient, partly calcified and and food availability). Below 17°C both adults and Table 1. Malacocoenosis from the site in Zieleniec <ul style="display: flex;"><li style="flex:1">Species </li><li style="flex:1">Ecological components </li></ul>hygrophileEuro-Sibe euryoecious euryoecious euryoecious woodland Zoogeographic composition Carychium minimum Cochlicopa lubrica Columella edentula Vertigo pusilla rian Holarctic Holarctic European Vertigo substriata Acanthinula aculeata Punctum pygmaeum Discus rotundatus Arion subfuscus N and E European W Palaearctic Holarctic woodland euryoecious euryoecious euryoecious woodland W and C European European Arion silvaticus C and NW European Holarctic Vitrina pellucida Eucobresia diaphana Vitrea diaphana euryoecious hygrophileC Europe hygrophileCarpathian-Alpine euryoecious woodland an Nesovitrea hammonis Oxychilus cellarius Limax cinereoniger Lehmannia marginata Deroceras leave Palaearctic W European S and W European European woodland woodland hygrophileHolarctic woodland Deroceras praecox Euconulus fulvus Cochlodina laminata Macrogastra badia Alinda biplicata W Carpathian-Sudetes <ul style="display: flex;"><li style="flex:1">Holarctic </li><li style="flex:1">euryoecious </li></ul><ul style="display: flex;"><li style="flex:1">woodland </li><li style="flex:1">European </li></ul><ul style="display: flex;"><li style="flex:1">woodland </li><li style="flex:1">E Alpine </li></ul><ul style="display: flex;"><li style="flex:1">woodland </li><li style="flex:1">C European </li></ul>Perforatella incarnata Trichia hispida <ul style="display: flex;"><li style="flex:1">woodland </li><li style="flex:1">C and SE European </li></ul><ul style="display: flex;"><li style="flex:1">European </li><li style="flex:1">euryoecious </li></ul>woodland Arianta arbustorum C and NW European 58 Tomasz K. Maltz, Beata M. Pokryszko Fig. 4. Ecological composition of the Zieleniec malacocoenosis Fig. 5. Zoogeographical composition of the Zieleniec malacocoenosis Fig. 6. Macrogastra badia. Variation in the number of apertural folds. Semi-diagrammatic Macrogastra badia (C. Pfeiffer, 1828) in Zieleniec 59 <ul style="display: flex;"><li style="flex:1">Table2. Macrogastra badia. Shell parameters of specimens </li><li style="flex:1">Table3. Macrogastra badia. Egg characters, egg number per </li></ul><ul style="display: flex;"><li style="flex:1">from the site in Zieleniec (n=39) </li><li style="flex:1">batch and incubation period </li></ul><ul style="display: flex;"><li style="flex:1">H</li><li style="flex:1">W</li><li style="flex:1">h</li><li style="flex:1">w</li><li style="flex:1">RangaenMe SD </li></ul>Range (mm) <ul style="display: flex;"><li style="flex:1">10.8–13.12 2.92–3.28 2.5–2.93 2.07–2.36 </li><li style="flex:1">Egg characters (n = 16) </li></ul>Major diameter (a) [mm] Minor diameter (b) [mm] a/b 1.39–1.61 1.32–1.45 1.00–1.13 1.51 1.39 1.08 0.089 0.039 0.044 Mean SD 11.88 0.496 N 3.11 0.08 folds 2.74 0.105 H/W 2.18 0.067 h/w Egg number per batch (n = 8) <ul style="display: flex;"><li style="flex:1">Range9.25–10.5 </li><li style="flex:1">0.5–4 </li><li style="flex:1">3.56–4.18 1.08–1.39 </li></ul><ul style="display: flex;"><li style="flex:1">1–3 </li><li style="flex:1">2.38 </li></ul>17.2 0.744 0.919 Mean SD 9.87 0.332 2.67 0.719 <ul style="display: flex;"><li style="flex:1">3.82 </li><li style="flex:1">1.25 </li></ul>0.058 Incubation period [days] (n = 10) 0.159 16–19 Fig. 7. Macrogastra badia. Proportion of various types of fold arrangement Fig. 8. Macrogastra badia: A – e ggsizeversus adult size , B – eggs, C – hatching, D – three-week-old juveniles, E – five-month-old juveniles. Scale bar 1 mm (Photo T. K. MALTZ) 60 Tomasz K. Maltz, Beata M. Pokryszko juveniles showed a considerable activity. At higher early in the morning. When the temperature extemperatures they remained hidden (moss, pieces of ceeded 20°C (July–August 2008), thesnails showed timber), immobile, retracted deep into their shells. no activity and soon most of them (70%) died. Sometimes single active specimens could be seen DISCUSSION The newly discovered locality of M. badia in Ziele- Arion silvaticus, Limax cinereoniger, Lehmannia niec, and the earlier Polish and Czech records (LOZEK marginata, Cochlodina laminata, Perforatella incarnata 1955, 1956, 1964, WIKTOR 1964, BRABENEC 1973, and Arianta arbustorum – all common and widely disHLAVÁÈ & HORSÁK 2002, HLAVÁÈ 2003), form a group tributed. The accompanying hygrophiles, recorded of localities which are remote and isolated from the from Zieleniec but not from Šumava, are Carychium main range of the species; they are located at an alti- minimum, Vitrea diaphana and Deroceras laeve. Also the <ul style="display: flex;"><li style="flex:1">tudeof 700 to morethan 900 m a.s.l. They aremainly </li><li style="flex:1">IN values show the small similarity between the </li></ul>natural deciduous and mixed forests with a consider- malacocoenoses. It can be supposed that M. badia is able proportion of beech and sycamore maple, with not a part of any definite malacocoenosis and its ocrotting logs and a rich herb layer (WIKTOR 1964, currence is determined solely by the environmental 2004b, HLAVÁÈ & HORSÁK 2002 and references cited factors. Theaccompanying malacofauna may repre - therein). The occurrence of the species seems to be sent either an original malacocoenosis which survived closely associated with the plant community of a com- in an unchanged habitat (sites in nature reserves in position similar to that in Zieleniec, identified by Šumava; HLAVÁÈ & HORSÁK 2002), or result from a ŒWIERKOSZ (2003) as Aceri-Fagetum and characteristic random colonisation of naturally regenerating huof the Alps. Such places, located at considerable alti- man-transformed habitats from adjacent areas (overtudes (800–900 m a.s.l.), provide adequate conditions: growing montane meadows in Zieleniec) (ŒWIERKOSZ a) lower vegetation-season temperature compared to 2003). A similar situation has been observed in the lower-situated areas, b) considerable humidity ensured caseof fauna accompanying another Alpine by the dense canopy, rotting timber and frequent pre- clausiliid, Charpentieria ornata, in Poland and the cipitation, and c) logs and largeamounts of litter providing the snails with shelter during the winter. Re- Czech republic (MALTZ 2009). The shell measurements of specimens from cently described localities in Šumava (HLAVÁÈ & Zieleniec do not depart from those reported by HORSÁK 2002, HLAVÁÈ 2003) are located at lower alti- LOZEK (1964), WIKTOR (1964) or HLAVÁÈ & HORSÁK tudes (720–750 m a.s.l.), but they are close to streams, (2002) (Table 4). The only essential difference is the on northern slopes with frequent thermal inversion. In number of apertural folds (0.5–4; according to LOZEK <ul style="display: flex;"><li style="flex:1">such conditions M. badia can livealso in partly open </li><li style="flex:1">1956, 1954 it is 1–2, according to KERNEY e t al. 1983 – </li></ul>habitats (Debrník stream valley on the Czech-German 1–3), and the number of 2–3 folds can be regarded as border), where abundant herbs and stones compen- typical of the discussed population. The arrangement <ul style="display: flex;"><li style="flex:1">sate for the absence of trees and dead timber. </li><li style="flex:1">and appearance of apertural lamellae conform to the </li></ul>Natural deciduous and mixed forests with condi- literature descriptions (LOZEK 1956, 1964, WIKTOR tions favourablefor M. badia are at present preserved 1964, 2004a i b, KERNEY et al. 1983). only as small and scattered fragments. Their gastropod faunas vary widely in their composition and it is thelifehistory of M. badia. Our laboratory observapractically impossibleto point to a charactreistic tions show that it is oviparous, likenineother native Published literature contains no information on malacocoenosis. The literature information is scanty clausiliids (MALTZ & SULIKOWSKA-DROZD 2008). Its and pertains only to recently described localities eggs are partly calcified, but – contrary to the situa(HLAVÁÈ & HORSÁK 2002 and literature cited therein, tion observed in the other studied native clausiliids – <ul style="display: flex;"><li style="flex:1">HLAVÁÈ 2003). Becauseof thecharacter of thehabitat </li><li style="flex:1">calcium carbonate in the egg envelopes, instead of </li></ul>it is not suprising that forest-dwelling species form a distinct crystals, forms a morecontinuous layer. The majority. Semilimax semilimax, S. kotulai, Malacolimax egg size is similar to those observed for Cochlodina tenellus, Clausilia cruciata and Macrogastra ventricosa orthostoma, Charpentieria ornata, Macrogastra latestriata, (HLAVÁÈ & HORSÁK 2002) are regarded as character- Clausilia dubia or C. pumila – species of similar shell <ul style="display: flex;"><li style="flex:1">istic of thegroup of localities in Šumava. Noneof </li><li style="flex:1">size(M ALTZ & SULIKOWSKA-DROZD 2008). However, </li></ul>these species has been found in Zieleniec though all theeggs of M. badia aremorespherical, compared to live in the Polish part of Central Sudetes (WIKTOR the eggs of these species, and their form is probably 1964, POKRYSZKO & MALTZ 2007, POKRYSZKO e t a.l determined by the more compact carbonate layer. 2008). Forest-dwelling species occurring both in The number of eggs per batch is similar to such num-

Macrogastra Badia (C. Pfeiffer

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