Of the Ship Rat in Lowland Forest 21
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DANIEL: DIET OF THE SHIP RAT IN LOWLAND FOREST 21 SEA SON A l DIE T 0 F THE S HIP RAT (R A T T US R. RAT T US) IN lOWLAND FOREST IN NEW ZEALAND M. J. DANIEL Ecology Division) DSIR, Lowei~Hutt ,.".; ~ SUMMARY: The seasonal diet of ship rats in a stand of lowland podocarp~rata~broadleaf forest in North Island, New Zealand, was studied from analysis of 173 stomachs, 46 fresh droppings and 10 feeding trials. Arthropods, particularly tree-wetas. (Order Orthoptera),. were the main foods in spring and summer, while drupes, berries and nuts predominated in autumn and winter. Birds were not an important food. Seasonal variations in the diet were related to the seasonal abundance of these foods in the forest. INTRODUCTION the distribution and abundance of geckos and skinks (Whitaker 1968) and possibly of tuatara The European ship, roof or bl~k rat (Rattus r. (S phenodon punctatus) on islands, and of land rattus L.) is one of three species of rats introduced snails (Paryphanta and Placostylus spp.), native and now established in New Zealand. The others frogs (Leiopelma spp.) and rare endemic arthro- are the Norway rat (Rattus n. norvegicus Berk.) f pods such as the gjant ground weta (Deinacrida and the Polynesian rat or kiore (Rattus e. exulans spp.) (Buller 1870). Peale). Ship rats probably arrived in New Zealand in the late eighteenth and early nineteenth cen- Beveridge (1964) also found that ship rats can turies and are now widespread in to'flTlS and forests consume almost the entire seed crop of podocarp , (Dieffenbach 1843, White 1895, Wodzicki 1950, trees such as rimu (Dacrydium cupressinum), . Watson 1953). matai (Podocarpus spicatus) and kahikatea (P. dacrydioideJ) and of hardwood species such as Apart from several short notes published in the hinau (Elaeocarpus dentatus). Ring-barking of late 1800s in which the species of rat mentioned is young Pinus radiata by ship rats has been reported often open to doubt (e.g. Wilson 1878, Kingsley in Chile (Riihm 1966)-but not so far in New 1895, Buller 1871), and general surveys (Wodzicki Zealand, although podocarp and other timber tree 1950, Watson 1953, 1959), the only recent papers seedljngs are destroyed by Norway rats (Beveridge on the ecology and spread of the ship rat in New and Daniel 1965). Zealand are those by Best (1969), Daniel (1972) and Atkinson (in press) . This paper describing the seasonal foods of ship rats in indigenou~ forest is part of a long-term Buller (1873), Stead (1936), Thomson (1922) study of rodent populations and their food supply and others have suggested that ship and Norway in lowland podocarp-rata-broadleaf forest in the rats are at least partly responsible for the marked Orongorongo VaIley. Population dynamics, home reduction in numbers of species and abundance of range and reproduction have been described endemic birds. Recently ship rats irrupted on Big (Daniel 1972). South Cape Island, off Stewart Island, and exter- minated four species or subspecies of rare birds on THE STUDY AREA the island (Blackburn 1965, Merton 1965), as well as probably extenninating the southern short-tailed The study area covered about 16 ha of forest \ bat (Mystacina tuberculata robusta) (Messrs D. V. near the DSIR Field Station (41021'S, 174°58'E) ,Merton and J. McIntosh, pers. comm.). However, in the Orongorongo Valley, Wellington province. \ Best (1969) found that birds were not an import- The locality and forest vegetation have been de- ; ant food of ship rats in long established rat popu- scribed (Daniel 1972). Briefly, the forest consists !lations jn t\,\'o South Island forests. Rats may affect of three main strata: a discontinuous upper stra- / 22 PROCEEDINGS OF .THE NEW ZRALAND ECOLOGICAL SOCIETY, VOL 20, 1973 turn of emergent northern rata (M etrosideros material was identified by' comparing the colour, Tobusta) and podoca~s, (mainly rimu, I~atai ,a~d ,texture and thickness of exocarp, peri-carp and .L ",';,' '",,' miro (P. ferrugineus) varying in height frofu aJ)out endocarp fragments with a reference collection of 25m to over 35m; a discontinuous middle stratum ripe and unripe berries, drupes, seeds and nuts at about 12m to 24m consisting of several species from the study area. For arthropods, fragments of of broadleaf trees-mainly hinau, pukatea (Laur- antennae, legs and exoskeletal material were com~ elia novaezelandiae) and rewarewa (K nightia ex- pared with those of a reference collection of arth~ celsa) ; and a continuous lower stratum from about ropods. Unidentified fragments were preserved in 3m to 12m composed of small trees and shrubs. 90% alcohol for later determination by experts. Epiphytes, particularly Astelia solandri, Collo- Three methods were used to quantify stomach spermum hastatum and Freycinetia banksii, Hanes contents: such as Ripogonum scandens, Metrosideros fulgens and Muehlenbeckia australis, and five species of (1) Items were recorded as present or tree ferns are conspicuous features of this forest. absent in each stomach and the percentage frequency of occurrence was calculated (the Mean annual rainfall at the Field Station over number of stomachs in V\'hich an iteni occurred an eight year period, 1953-1960, was 2443mm expressed as a percentage of the total number range 2032 - 2743mm). Mean summer and winter of stomachs examined). temperatures over the four years 1966.69 were 15.5°C and 6.7°C respectively. Mean winter mini- (2) The number of occurrences of each mum was 2.8°C and mean summer maximum was food item was expressed as a percentage of 19.4°C. Prevailing westerly winds create turbulent the total of occurrences of all items, following conditions on the valley floor, and on the exposed Hynes (1950) and Rudge (1968). ridges gusts in excess of 160 km/hr (100 mph) are (3) A visual estimate to the nearest 10% frequent. was made of the relative volumes occupied by plant and animal matter in the contents of METHODS each stomach. L Collection of Rats Hansson (1970) discussed the usefulness and The total sample of 173 rats comprised 31 snap- limitations of these three methods. No attempt trapped on three lines from November 1966 to was made to weigh the stomachs or food items as October 1967, 125 -rats snap-trapped on and around done by Kami (1966) and Fall, Medina and the live-trapping study area from August 1968 to Jackson (1971). July 1969 and 17 rats killed during live-trapping To supplement the stomach analyses, 46 fresh studies (see Daniel 1972). droppings were collected during live-trapping from the traps and from live animals: 12 in spring 2. Stomach and Faecal Analysis (September to November), 10 in summer (Decem- Stomachs were removed from freshly killed rats ber to February), 12 in autumn (March to May) . and preserved in 90% alcohol. For analysis the and 12 in winter (June to August). The droppings . contents were placed in a petri dish and examined for each season were grouped, mixed with water in under a dissecting microscope. Fifty-three stomachs a petri dish and examined under a dissecting micro- ' out of a total of 173 (31 %) contained unidenti- scope. Although more finely comminuted than fied amorphous starchy material probably repre- many of the stomach samples, the contents of the senting kernels of hinau, rniro or nikau (Rhopalo- droppings gave a rough indication of the seasonal stylis sapida) nuts. Many food items could be frequency of plant and insect foods, and some readily identified,- e.g. the speckled femurs of tree plants and insects could be identified from smiH wetas (Hemideina thoracica), the spotted endo- fragments. The seasonal changes in plant and' carp of supplejack (Ripogonum scan dens ) berries animal foods followed ,the same trends as those; 'and the green exocarp of hinau drupes. Plant from stomach contents. DANIEL: DIET OF THE SUIl' RAT IN LOWLAND FOREST 23 TABLE 1. Analysis of 173 ship rat stomachs showing number of occurrences of each food item, its percentage frequency of occurrence and contribution to the diet. SPRING SUMMER AUTUMN WINTER ALL SEASONS % % % % % % % % % % No. Occur. Diet No. Occur- Diet No. Occur. Diet No. Occur- Diet No. Occur. Diet ence ence ence ence ence - - PLANT MATTER Unid. pericarp 10 24 16 3 7 4 24 55 36 16 43 30 53 32 20 Elaeocarpus dentatus 0 0 0 0 0 0 2 5 3 7 19 13 9 5 3 Hedycarya arborca 2 5 3 7 16 9 1 2 1 3 8 6 13 8 5 Rhipogonum scandens 3 7 5 0 0 0 6 14 9 5 14 9 14 8 5 Rhopalostylis sapida 0 0 0 0 0 0 2 5 3 1 3 2 3 2 1 Coprosma spp. 0 0 0 0 0 0 0 0 0 2 5 4 2 1 1 Schefflera digitata 1 2 2 0 0 0 0 0 0 1 3 2 2 1 1 Corynocarpus laevigatus 0 0 0 1 2 1 0 0 0 0 0 0 1 1 I Macropiper excelsum 0 0 0 3 7 4 8 18 12 2 5 4 13 8 5 Green leaves'*' 1 1 27 18 10 22 13 4 9 6 2 5 4 27 16 10 Fungi 0 0 0 0 0 0 0 0 0 4 1 1 7 4 2 2 STOMACHS WITH PLANT MATTER 27 64 - U 51 - 40 89 - 34 87 -- 125 72 - ANIMAL MATTER Unid. Arthropods 4 10 7 6 13 8 4 9 6 1 3 -? 15 9 6 Orthoptera H emideina thoracica 22 54 36 25 56 33 15 34 22 5 14 9 67 40 26 Unid. Orthoptera 0 0 0 0 0 0 1 2 1 0 0 0 1 1 1 Lepidoptera adults 1 2 2 1 2 1 0 0 0 0 0 0 2 1 1 larvae 2 5 3 0 0 0 0 0 0 2 5 4 4 2 2 Phasmatodea Phasmatoda 2 5 3 2 4 3 0 0 0 1 3 2 5 3 2 Coleoptera 2 2 3 5 11 7 0 0 0 0 0 0 7 4 3 Hemiptera CicadeIlidae 0 0 0 8 18 1 1 0 0 0 0 0 0 8 5 3 Diptera pupae 0 0 0 2 4 3 0 0 0 1 3 2 3 2 1 Araneae Araneidae 0 0 0 0 0 0 0 0 0 1 3 2 1 1 1 Hymenoptera Formicidae 1 2 2 0 0 0 0 0 0 0 0 0 1 1 1 Mollusca Wainuia urnula 0 0 0 2 4 3 0 0 0 0 0 0 2 1 1 Aves U nid.