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Final Comprehensive Conservation Plan - U.S. Fish and Wildlife ...

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Hakalau Forest National <strong>Wildlife</strong> Refuge<br />

<strong>Comprehensive</strong> <strong>Conservation</strong> <strong>Plan</strong><br />

80 days <strong>and</strong> begin mating 30 days after hatching. Both larvae <strong>and</strong> adult koa bugs emit odorous<br />

defense compounds when disturbed (Johnson et al. 2005).<br />

The koa bug has no known natural predators; however, nonnative spiders <strong>and</strong> ants are known to<br />

parasitize C. blackburniae eggs, nymphs, <strong>and</strong> adults. The big-headed ant chews koa bug eggs <strong>and</strong><br />

Western yellowjacket wasps are predators of young (Johnson et al. 2005). Other species known to<br />

kill koa bugs include Anastatus sp., Acroclisoides sp., <strong>and</strong> birds. In addition, two biocontrol agents<br />

(Trissolcus basalis <strong>and</strong> Trichopoda pilipes) that were introduced to control the southern green stink<br />

bug are able to locate <strong>and</strong> develop on C. blackburniae. Although these species have a relatively<br />

minor impact on C. blackburniae overall, substantial impacts may occur at individual sites (Johnson<br />

et al. 2005).<br />

The koa bug has also been impacted by the reduced abundance <strong>and</strong> distribution of its host species as<br />

a result of agricultural activities <strong>and</strong> urban development. Because these plants typically fruit at the<br />

same time, locating host plants suitable to lay eggs may require C. blackburniae to disperse over long<br />

distances. Studies have shown that koa bug mortality due to dispersal accounted for about 50 percent<br />

mortality in all individuals studied (Johnson et al. 2005).<br />

4.8.3 Cave Invertebrates<br />

Until the 1970s, it was assumed that the young <strong>and</strong> geologically unique Hawaiian Isl<strong>and</strong>s did not<br />

support an abundance of cave-adapted fauna. However, Hawaiian caves have been found to support a<br />

diverse array of rare <strong>and</strong> highly specialized invertebrates (Howarth 1972, 1983, 1991). These<br />

cavernicoles can be classified into three categories: (1) troglobites, which are obligate cave species;<br />

(2) troglophyles, which can live in caves or other cave-like (moist cool dark) habitats; <strong>and</strong> (3)<br />

trogloxenes, which can be found in caves, but do not live their entire life in caves (Howarth 1973,<br />

1983).<br />

The Service recognizes one species of cave invertebrate on the Isl<strong>and</strong> of Hawai„i, the troglobitic<br />

cixid leafhopper Oliarus polyphemus, as a species of concern. Troglobites are only able to survive in<br />

cave ecosystems. These species are characterized by a number of anatomical <strong>and</strong> physiological<br />

adaptations to cave life (Barr 1968, Christiansen 1982, Holsinger 1994, Culver et al. 1995).<br />

Troglobites tend to lose their pigmentation, eyes, <strong>and</strong> hard exoskeletons. Additionally, troglobites<br />

have elongated appendages <strong>and</strong> sensory structures with long hairs, lengthened life span, <strong>and</strong> modified<br />

life-history patterns. As a result of their nutrient-poor environments, the life cycle of many<br />

troglobites is characterized by delayed reproduction, increased longevity, lower total egg production,<br />

<strong>and</strong> production of larger eggs (Culver 1982).<br />

Obligate cave species have narrow <strong>and</strong> specific ecological requirements. Such requirements include<br />

high relative humidity, stable temperatures, <strong>and</strong> a preference for high CO2 levels (Howarth <strong>and</strong> Stone<br />

1990). They do not acclimate well to rapid changes in their physical, biological, or chemical<br />

environment (Barr 1968, Culver 1982). Troglobites also rely on energy <strong>and</strong> nutrient input from the<br />

surface. These invertebrates primarily feed on „ōhi„a roots or other plant roots that penetrate the lava<br />

tube roof (Howarth 1973; 1983, Howarth et al. 2007). Additional nutrient sources into caves include<br />

plant detritus washed in by surface waters, organisms that enter caves under their own power, <strong>and</strong><br />

guano from bats, rats, <strong>and</strong> mice. Thus, cave systems can be strongly influenced by the surface<br />

ecosystem (Barr 1968, Culver 1982).<br />

4-48 Chapter 4. Refuge Biology <strong>and</strong> Habitats

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